Академический Документы
Профессиональный Документы
Культура Документы
by
AL JO S FA R JON
Volum e I
LEIDEN-BOSTON
2017
This book is printed on acid-free paper.
Library of Congress Cataloging-in-Publication Data
The Library of Congress Cataloging-in-Publication Data is available from the Publisher.
front cover:
Abies fabri young seed cones
back cover:
top left: Dacrycarpus kinabaluensis seed cones
bottom right: Picea likiangensis young seed cones
Foreword 7
Preface 9 5
Appendix 1071
Glossary 1073
References 1093
With the Handbook of the Worlds Conifers, its because it gives us the keys to designing effective
author Aljos Farjon has given us a wonderful aid to conservation measures for each endangered species
learn almost all there is to know about this ancient and also defines the scope and limitations of where
lineage of woody plants. In true Handbook tradi- one could introduce a species successfully in tropical
tion all 614 species of extant conifers are concisely or temperate horticulture. 7
described, with user-friendly identification keys to
families, genera and species and invaluable addi- With his lifelong dedication to the study of the tax-
tional information on uses, conservation status, and onomy of conifers at Utrecht, Oxford and Kew, and
much more. Unlike earlier books on conifers that are his strong involvement in the conservation of many
heavily skewed towards temperate conifers from the red-listed conifer species throughout the world,
northern hemisphere, the present Handbook does Aljos Farjon is arguably the only plant scientist in
equal justice to the 200 tropical species and 415 tem- the world to complete the colossal task of produc-
perate ones; this attention to tropical species makes ing this comprehensive handbook single-handedly
the Handbook truly unique. and authoritatively. I hope and am confident that it
will give as much pleasure and satisfaction to a wide
In the very informative introduction the evolution, audience of tree enthusiasts, foresters, botanists,
classification, ecology, biology, economic uses, and ecologists and conservationists as it has given me.
conservation status of the conifers are discussed.
This handbook thus combines the virtues of a com- Pieter Baas
prehensive taxonomic monograph of the conifers, Emeritus Professor of Systematic Botany, Leiden
with that of a true vademecum on the morphologi- University
cal attributes, uses, and conservation of all species Former Director of the National Herbarium of the
of the order. The information on the ecology of each Netherlands
species in its natural habitat is particularly welcome,
P R E FAC E
Many books have been written about conifers dur- are paragraphs on ecology, uses and conservation
ing the 19th and 20th centuries, the majority of these with all species accounts. This handbook does not
with emphasis on horticulture in Europe. Those that describe or illustrate conifers that are only known
are mostly compilations of species and their culti- in cultivation. With some 15,000 conifer cultivars
vars found to be grown in gardens and parks, with known, no comprehensive coverage could have 9
descriptions, illustrations and information about been achieved. Instead, a separate volume now cov-
their cultivation, are known as handbooks or manu- ers these, authored by Aris Auders in Latvia and
als. A conifer handbook as traditionally compiled is Derek Spicer in England, entitled Conifers A
therefore a comprehensive guide to the conifers that Comprehensive Guide to the Conifer World, pub-
are grown or could be grown in cultivation in a cer- lished under the auspices of the Royal Horticultural
tain part of the world and includes, besides species Society. It follows the taxonomy in this Handbook of
and botanical varieties, cultivated varieties or forms the Worlds Conifers for families, genera and species
(since 1953 universally known as cultivars). All such and these two handbooks are therefore complemen-
works, even the most comprehensive, emphasise tary and can (and should) be used side-by-side.
conifers adapted to a cool temperate climate as expe- The need for a comprehensive and modern hand-
rienced in Europe. Several have made this explicit book concentrating on the species of conifers is clear.
in the title of the book, but even those that have Awareness of the obligation to manage and pro-
not done this invariably treat hardy conifers more tect the biological resources of this planet and use
fully than other species, which are at best only men- these resources sustainably has created a demand
tioned briefly. The scope, content and presentation for accurate, up-to-date information. Conifers occur
are aimed at the horti- or silvicultural users. Some of worldwide and are major or minor components of
the better-known conifer handbooks were reviewed nearly all major forest and woodland types, as well as
in Chapter 1 of my Monograph of Cupressaceae and many timber plantations and numerous gardens and
Sciadopitys (Farjon, 2005a). Most conifer handbooks parks. They are of high economic value, but also play
are now out-of-print, but some new books have major roles in the ecology of many natural and semi-
recently been added to the bibliography of conifers, natural ecosystems. They are an integral part of many
while others are being compiled. The emphasis on landscaped estates and parks from the subtropics of
the temperate conifers in cultivation has remained Australia and New Zealand to the temperate regions
strong in these works, as far as I know. of Europe and North America. This handbook of
The present Handbook is very different from conifers has been compiled to address this need for
these predecessors. There is no particular emphasis relevant information and should therefore be useful
on conifers that can be grown in Europe. Of the 614 to botanists, conservationists, dendrologists, ecolo-
species known in the world at present, about 200 gists, foresters, horticulturists, land resource manag-
occur in the tropics. They receive here equal cover ers and, I should add, all those with an interest in
with the species that grow in temperate climates. In the trees and forests of the world. Knowledge is not
this respect, the present book more closely resem- only useful, it is a pleasure that enhances the quality
bles a taxonomic monograph, as it includes all the of life.
currently known and accepted taxa, from the ranks The introductory chapters have deliberately been
of family down to variety and forma. However, it is kept brief, often merely touching on interesting
not a monograph purely for taxonomists. Its con- aspects of conifer biology, evolution and taxonomy,
tent aims at a much wider audience. To achieve this, as well as ecology, economics and conservation. For
strictly taxonomic information has been limited to a recently published full introduction I refer to my
what is more or less essential, while other infor- book A Natural History of Conifers (Farjon, 2008),
mation has been included that may not even have where all these issues are treated in much more
been considered for a taxonomic monograph. There detail. After these short chapters, the book moves to
the families, genera and species and becomes the its homotypic synonyms) or begin on a new line.
handbook referred to above. For the general user it Where relevant, the (aberrant) status of synonyms
may be useful to outline and explain the conventions under the Botanical Code (ICBN) is given after
and format followed throughout for all genera and the citation of the place of publication. Types (i.e.
species. To identify a completely unknown conifer, herbarium specimens designated to fix the appli-
one should start with the key to families on p. xxx; cation of a botanical name) are only given here for
then the key to genera under the family determined, accepted names, including their synonyms if based
and the key to species under the genus arrived at. on the same type. Types of genera are cited as names
Genera with numbers of species greater than 1520 of species. The opportunity to designate types where
have been divided, formally or informally, and keys it was appropriate and possible has been taken; but
10 to these groups are first to be consulted. Under spe- in several cases typification is complicated and then
cies may follow subspecies and/or varieties; these it has been left to future research. This is what is
are usually not included in keys. Not more than six meant with the phrase type not designated under
infraspecific taxa are accepted under any one spe- several species. Type specimens are deposited in
cies in this book, and their descriptions are brief institutional herbaria, abbreviated by an acronym
and diagnostic. These descriptions should primar- and these can be found at www.sciweb.nybg.org/
ily be compared with the relevant species descrip- science2/IndexHerbariorum.asp.
tion and the character states mentioned therein, and Under species, subspecies, varieties and forms
less with each other. No attempt has been made to may be recognized. While these ranks are hierar-
create varieties within subspecies and, unless such chical in that order and species have been subdi-
ranking has been established by other authors and vided in that way, I am no advocate of this. A single
is followed here, the two ranks are treated as equal. rank under species would suffice in botany as well
In the taxonomic literature, the use of either sub- as it does in zoology. Leaving the rare use of forms
spieces or variety appears to derive more often from aside as a special case, I am agnostic as to whether
tradition than from explicit taxonomic concepts. I this should be the rank of subspecies or variety. In
believe that botany would be well advised to follow botanical taxonomy, we have an unavoidable his-
zoological practice and use only a single infraspe- torical legacy: both ranks have been used in the
cific rank in future. After using the keys, reading the literature, often for the same taxon. In some coun-
description of a species is recommended as a check tries one rank was preferred over the other. In this
to find out if the determination is correct. Also, as it Handbook, therefore, I have used what ranks were
is almost impossible to construct keys that use any available in validly published names under species
and every feature of a specimen in hand, the reading where infraspecific taxa merited recognition; under
of the description helps with the identification. An a few species I have accepted two ranks if these
extensive glossary is provided to help explain much appear to be widely used. Unfortunately, the same
of the terminology used in this handbook. notion of one rank under species has induced the
Botanical names are given in full as in a taxonomic publication of numerous new combinations, some
monograph, i.e. with (abbreviated) author name(s), have changed subspecies to variety and others vice-
abbreviated reference to the place of first publication, versa depending on preferences. I have not seen fit
synonymy and type citation. The accepted names are to increase the number of pages by including all of
followed by synonyms if these apply. Among syn- these mostly superfluous name changes, nor by add-
onyms, basionyms are always cited as types refer to ing new combinations if none existed in the desired
these; other synonyms are cited selectively. In most rank. If infraspecific taxa are recognized, the spe-
cases, synonyms no longer in use as accepted names cies name in the inclusive sense is given first (with
have been omitted. Most recently published names authority and place of publication) and the descrip-
have been included, but not all, for instance new tion is similarly inclusive, i.e. it encompasses all
names resulting from a mere change of rank, such subspecies and/or varieties. Synonyms and types
as variety to subspecies. When several synonyms are are given with the latter and the short descriptions
cited, homotypic synonyms are grouped together in given there are exclusive. Botanical descriptions
one continuous line; heterotypic synonyms are sepa- immediately follow the names of taxa and are not
rated by a blank line from the accepted name (and headed, but all other information is given under its
specific head (in bold type face) for clarity and ease entered in a database, the TDWG codes linked to
of reference. The descriptions of species follow the names can provide easy and quick listings of spe-
conventional sequence for descriptions of trees; i.e. cies per country, state or province. This information
from size and habit to trunk, bark, branches, foliage, is available for conifers at www.catalogueoflife.org/
leaf details, male and female reproductive struc- search.php.
tures and ending with seeds. Despite the relative Ecology describes the habitat of a species and its
detail in some descriptions, it should therefore not altitudinal range (in meters above sea level: m a.s.l.)
be difficult to find particular parts and I have there- is given here. For many species, especially in the
fore not marked these with a different font or type. tropics, their habitat remains poorly documented,
The botanical names of species are followed by the while much has been written on many species in
etymology of the accepted name and by vernacular the temperate northern hemisphere. Some infor- 11
names. Etymology is also explained for genus names mation could be assembled from notes on herbar-
but not for subspecies and varieties and only occa- ium specimen labels and has been recorded in the
sionally vernacular names are mentioned with the conifer specimen database available at dps.plants.
latter two. The vernacular names do not provide a ox.ac.uk/bol//home/default.aspx under Online
comprehensive listing, especially not for species with Groups. As indicated above, a balanced approach
a very large natural distribution, covering an area in has meant that not all that is known can be men-
which many languages may be spoken. Vernacular tioned here and for such species the information on
names may not be specific; if referring to the genus ecology (and other subjects) is indicative rather than
rather than the species and if more than one species comprehensive. Conservation is usually concerned
occur in the area, such names are not listed. Some with species survival issues as defined by the World
published vernacular names seem contrived; if these Conservation Unions (IUCN) Species Survival
are not actually in use locally or regionally, they Commission (SSC). Information was gathered and
are not cited here. Taxonomic notes may follow the the conservation status of taxa (species to varieties)
descriptions; they are usually comments on different has been assessed by members of a Specialist Group
taxonomic treatments or notes on typification of the (SG) resorting under the SSC. In the case of conifers
species or mention similarities of taxa. In the con- this is the Conifer Specialist Group (CSG) which is
cise format necessary in this Handbook, comments currently chaired by this author. Almost all conifers
and discussions are limited to the essential and are have been assessed under IUCN Red List criteria,
kept brief. Not every name placed in synonymy can the resulting categories of threat are given under
be discussed here; the reader can rest assured that the heading IUCN. For the abbreviated categories
due scientific consideration has been given, some of of threat see the introductory chapter Conservation;
which has been published elsewhere. Distribution for definitions and decoding of the criteria used
of taxa is given in two formats and these are often visit www.iucnredlist.org/static/categories_criteria.
complementary. The first is descriptive, mention- The listings given here are as in the 2008 IUCN Red
ing regions as well as countries, provinces, areas and List of Threatened Species [www.iucnredlist.org]
localities, as appropriate. The second format uses the including the criteria of either version 2.3 (1994)
TDWG codes (Brummitt, 2001). These codes pro- or 3.1 (2001) as applied. Uses mentions commer-
vide geographical information at three levels defined cial as well as other human uses of species. The two
by geographically and politically delimited areas, e.g. main categories are wood (timber) and its applica-
42 = Malesia (a geographical region) BOR = Borneo tions and horticulture, but other uses, including
(an island) and SB = Sabah (a politically delimited traditional, have been mentioned when known.
area): 42 BOR-SB means therefore that the species As with ecology, much more is known about uses
occurs (also) in Sabah. The meaning of these codes of species in the temperate zones than in the trop-
can be found at www.catalogueoflife.org/search.php. ics. Often, timber is traded under a generic name
For species with large ranges the TDWG codes pro- which may even include different genera, as in the
vide more detailed information than the descriptive family Podocarpaceae, which makes distinctions of
statement, but for species with very limited distribu- use among species extremely difficult. Horticultural
tion the latter is often more precise because TDWG use is also unevenly distributed along similar lines of
codes often do not go down to such detail. When geographical demarcation, but uses in the subtropics
and tropics are picking up and therefore mentioned had to be raised for this project, and when money
when known. This book does not include cultivars was being pledged, I had to find a registered char-
but if such exist under a species, that information ity to administer these funds. This was generously
is given. Plantation in a forestry context is usually undertaken by the Linnean Society of London and
mentioned with the uses of timber; for several spe- I am grateful for the trust the Society has placed
cies plantations have become the main source of that in one of their Fellows to fulfil the commitments
commodity. thereby generated. The Treasurer of the Society, Prof.
The glossary in this Handbook is the most inclu- Grenville Lucas, guided me through this whole pro-
sive I have yet compiled. I have made use of several cess in his capable and incomparable manner and I
published in my earlier works, but added numer- am most indebted to him for this generous support.
12 ous entries specifically for this book. In particular, Most importantly, my gratitude goes to all the gener-
I have attempted to explain terms used by plant tax- ous donors to this project, who together have made
onomists, because this book is meant to be used and this book possible. I have permission to name them
consulted by many others, amateur as well as profes- here and shall do so in alphabetical order. They are
sional. I hope it will be found useful, but I am aware the Arboricultural Association (UK), Mr. Lawrence
of the difficulty of finding a balance between explain- Banks (UK), The Dendrology Charitable Company
ing commonly understood terms and thereby under- and the International Dendrology Society, Dr. Barry
estimating my readers and using unexplained terms Denyer-Green (UK), the Lord Devonport Charitable
for which one might need to consult a large diction- Trust (UK), Mrs. Francine von Finck (Germany),
ary. Finally, I have only included botanical names of Mrs. Arabella Killander (UK), the Samuel Storey
conifers in the index. Numerous other species names Family Charitable Trust (UK) and the Stanley Smith
occur in the texts on the ecology of conifer species, Horticultural Trust (UK). This Handbook is the syn-
but giving information about these other species thetic product of 25 years of research into conifers
is obviously beyond the scope of this Handbook. by its author. To give detailed acknowledgement
Vernacular names, while cited under many species, to all persons and organisations who have assisted
are incomplete (especially in the tropics), use several me in that research would be next to impossible.
languages besides or in lieu of English, and are often Research is often a collaborative effort, even if the
ambiguous. The reader is therefore encouraged to resulting published output appears to have a single
use (and learn) the botanical names of species. Both author. My thanks are therefore here expressed to
in the library and on the Internet, nearly all infor- all I have been in touch with on conifers from time
mation is available under these identifiers. Because to time, on field trips around the world as well as at
the information in this book is arranged by species, my and their various home institutes during those
finding the species (they are arranged in alphabeti- years. Dr. Christopher N. Page reviewed the texts on
cal order as well as indexed) is all that is needed to Podocarpaceae and the smaller families and his sup-
obtain all the information about them given in this portive comments were much appreciated. Michael
Handbook. Frankis commented on common names and cor-
The compilation and writing of the Handbook of rected errors therein. The entire text was copy-edited
the Worlds Conifers has been a task which started by Dr. Hans Kruijer, whose helpful corrections much
some time after my official retirement from the Royal improved consistency and clarity of presentation.
Botanic Gardens, Kew in 2006. Certain conditions Prof. Pieter Baas wrote a thoughtful foreword, for
had to be met before I could seriously embark on which I am most grateful. Finally, coming closer to
this project, because it required access to resources, the end product now before you, I wish to thank
to enable me to devote full working time to it. I the publisher, Brill and their helpful staff (especially
was very fortunate that the Royal Botanic Gardens Michiel Thijssen, Sabine Steenbeek and Frits Fritschy)
appointed me an Honorary Research Associate after for their professional work and helpful patience with
my retirement, to be based in the Herbarium where the author. They have turned a huge and complicated
I had worked before (now the Herbarium, Library, manuscript into a beautiful book.
Arts & Archives Department). It was there that much
of the work on this book was done and I am grate- Aljos Farjon, FLS
ful to RBG Kew for this invaluable support. Funds [Kew, 27 November 2009]
P R E FAC E T O T H E SE C O N D
E D I T IO N
Since the publication of this Handbook in 2010 on distribution and conservation status of spe-
rapid and substantial developments in conifer cies became available and had to be incorporated
research and publication have occurred. Those in the new edition. Proposed taxonomic changes
that have caused many necessary corrections and have been treated cautiously but those considered 13
amendments to the first edition are connected unavoidable have been presented in an Appendix
with two projects led by the author; the compila- so as not to disrupt the alphabetical sequence
tion of An Atlas of the Worlds Conifers published adopted in the Handbook.
by Brill in 2013 and the reassessment for the IUCN
Red List of all conifers in 20102013. As a conse- AF
quence, new and often more detailed information [Kew, 30 November 2016]
T H E C O N I F E R S O F T H E WO R L D, A N I N T R O DU C T IO N
To most of us who live in the temperate zone of the But, such criteria of appeal to biologists aside, the
northern hemisphere, conifers are among the most sheer number and volume of conifer trees growing
familiar trees and indeed an everyday sight. Not in the temperate north makes them of prime eco
only do they naturally occupy vast areas of north nomic and ecological importance throughout the
ern lands such as Canada, Scandinavia and Russia, entire world. With an estimated value of $100 bil 15
as well as many mountain regions further south, we lion per annum, wood products stand foremost and
have also extensively planted them in forestry while conspicuous at the top of the economic shopping list,
gardens and parks of even moderate size are hardly but especially at more southern latitudes other prod
conceivable without them. Far less widely known ucts, e.g. resins and derivatives, and even seeds for
are the conifers of the southern hemisphere and still food, are also economically important. And then, as
fewer people will associate conifers with the tropics. already indicated, there is the significant contribu
True, neither of these major parts of the world have tion conifers have made to horticulture as ornamen
extensive conifer forests similar to those in the north; tal trees, a trade which has given rise to numerous
conifers in these latitudes are more often than not cultivars some of which are among the most popu
a relatively minor component of forests, often con lar shrubs and trees planted almost the world over
fined to rather inaccessible areas, or inconspicuously in gardens and parks. I venture the statement that,
mixed with more numerous angiosperm trees. But if although being a relatively small group of species of
we were to regard diversity rather than sheer quan woody plants there are about as many species of oak
tity as a criterion, the north would lose its primacy (Quercus) and twice as many species of wattle (Acacia)
as the realm of conifers to more southern latitudes. conifers far exceed any other group of woody plant
In the present Handbook of the Worlds Conifers I species in economic and ecological importance. We
have recognized 614 species in 70 genera belonging would have to look at the major food crop plants to
to 8 families. Of the genera, 35 are restricted to the find species that generate more income than some of
northern hemisphere, 25 occur only south of the the pines and spruces. But, as with cereals and beans,
equator and 10 are found both north and south of only a tiny minority of species diversity within coni
that line. Since, with the exception of Podocarpus, fers is accountable for this prominence today, while
which is largely tropical, most large genera occur in the majority shares in the fame only by merit of kin
the northern hemisphere, that half of the globe (with ship. Having said this, the question may be asked,
by far the greater land mass) has more species than what constitutes this kinship in conifers? In other
the southern hemisphere. But in the northern hemi words, what are conifers? It is not as straightforward
sphere, diversity of species, especially in the larger a question to answer as it may seem.
genera, increases markedly towards the equator, with In the early days of systematic botany, the coni
for instance only 9 species of Pinus in Canada and 43 fers were often thought of as a natural family of
in Mexico. More or less limited regions with more plants, in a similar fashion as botanists referred to
than 40 species or 50 taxa (species and lower ranks) e.g. Rosaceae or Compositae and, indeed, the name
defined as centres of diversity for conifers (Farjon Coniferae prevailed well into the early 20th century.
& Page, 1999) are now eight in number, seven are Are conifers a family of plants, or at a rank higher,
situated on or north of the equator and one south an order Coniferales? Modern classifications classify
of it, but all except Japan are at latitudes below 40 groups of organisms in monophyletic groups, i.e.
and two, Borneo and New Caledonia, are entirely in organisms or groups of organisms that in evolution
the tropics. That last island is the most diverse and ary history share a common ancestor (usually an
remarkable conifer centre of all; covering an area the ancestral species). Monophyletic groups may include
size of Wales, it has 43 species and all are endemic to more recently formed monophyletic groups and be
the island. long to other monophyletic groups of more distant
pasts. We must discuss the term conifer in this novelty of having ovules and seeds enclosed in cap
context of monophyly to answer these questions. sules, angiosperms form a natural group. The term
No such discussion makes any sense without look gymnosperm, on the other hand, does not specify
ing at the origin of conifers as (imperfectly) revealed a natural group by any modern criterion. Conifers
in the fossil record. The fossil story is a complicated are characterized by naked ovules, for sure, and are
tale which can only be very summarily related here, thus gymnosperms, but they share this one feature
but I shall nevertheless try to answer these ques of early seed plants with other groups of primitive
tions; for more on this topic, see my book A natural seed plants, e.g. Ginkgo and cycads. Taxa are defined
history of conifers (Farjon, 2008). The word conifer, by evolutionary novelties. It is, however, often dif
meaning cone-bearer, suggests that plants bearing ficult to determine which characters represent
16 cones could be called conifers. Cones are reproduc such evolutionary novelties. Taxa, therefore, need
tive organs consisting of fertile scales (simple cones) to be defined by diverse and structurally unrelated
or sterile and fertile scales (compound cones) that characters to increase the likelihood of represent
are spirally arranged at a central axis; the fertile ing natural groups and for conifers several of such
scales of a cone contain either pollen sacs or ovules. characters can be identified. Farjon (2008) charac
However, although the majority of conifers are char terized conifers as shrubs or trees with secondary
acterized by having cones, it is problematic to define wood build of tracheids with large bordered pits in
the conifers as plants bearing cones. Even if the term their walls and narrow rays. The leaves are simple
cone would be restricted to the ovulate organ of its and single or parallel veined. Resin is produced in
description, a definition of conifers based on that the wood or in the leaves and is conducted through
organ is problematic, because we will then find that, resin canals. The reproductive organs are separated
among the gymnosperms, pines are considered coni into male and female, with male cones (pollen cones)
fers but cycads are not. Conversely, Cephalotaxus, simple and female cones (seed cones) compound
with a mature ovulate organ reduced to a single or reduced. Conifers have only one copy of a large
seed is still a conifer, but Ginkgo, seemingly similar inverted repeat in the chloroplast DNA, whereas all
in that respect, is not (despite its inclusion in many other plants studied so far have two copies. If the
books on conifers). The key to the difference here is DNA observation holds true, it may well turn out to
partly revealed in the ontogeny of both taxa, but it be the single character state that is unique to coni
ultimately has to do with phylogeny, i.e. how these fers. But the group is exclusively circumscribed by
taxa are related in an evolutionary sense. this plus the other characters, each not unique, but
in combination only present in conifers. Such char
The term gymnosperm has fallen, and we need to acters are sometimes called traits. The seeds of cer
explain what is meant by it first. In the course of the tain coniferous taxa occurring in the Carboniferous,
early evolution of land plants, adaptations to water when the earliest seed-bearing plants became appar
stress i.e. dry conditions gradually led to the ori ent, tended to be borne on appendages axillary to
gin of the seed habit, with the evolution of ovules leaves. Reduction of these appendages in size and/
with an envelope (integument) to protect the vulner or number, planation, movement of ovules towards
able gametophyte phase from desiccation, coupled the base of the appendage and transformation of
with devices to catch pollen in a suitable micro-envi the subtending leaf to a bract as well as reduction
ronment and to conduct the growing pollen tube of the fertile shoot to a determinate axis led to the
to the megagametophyte for fertilization. As these formation of the conifer cone. By these transforma
ovules were borne variously on branching systems, tions the female reproductive organs were aggre
leaves or other appendages, but otherwise exposed gated in compound cones, which later again became
to air and not enclosed in a capsule (ovary), like in reduced in various ways but the homology is often
angiosperms, they were naked as denoted by the still discernible in ontogeny. The male reproductive
term gymnosperm which is thus merely a denomi organs (pollen cones) remained, or became early
nator for a less advanced mode of seed habit. All on, a simple axis bearing microsporophylls with
seed plants were by definition gymnosperms until dehiscent microsporangia (pollen sacs). Pollen cone
the angiosperm seed habit evolved from a gym morphology is rather uniform throughout the coni
nospermous seed. By sharing the evolutionary fers. All these (and other) transformations are well
documented in the fossil record. They can generally exception of the Himalayan region which had a dif
be understood as adaptations to increasing arid ferent geological history), but rich in conifer fossils
ity and seasonality with cold winters, which indeed and those of other gymnosperms. On its long jour
marked the end of the Carboniferous and beginning ney from southern latitudes as a disconnected chunk
of the Permian in large sections of the superconti of Gondwana, going through the hot tropics and
nent Pangea. Out of perhaps a single ancestor (but one or two arid zones it must have lost them all. On
which one?) plants we now call conifers evolved, the collision with Asia, thrusting up the Himalayas,
spread across much of the ancient supercontinent, northern conifers spread southward to occupy the
were dominant almost everywhere when this was mountains. India has but a single southern coni
divided by the Thetys Ocean into Gondwana and fer today: Nageia wallichiana, which is in fact an
Laurasia, and prevailed even when these two con Indo-Malesian element spread from Southeast Asia. 17
tinents broke up and the present continents started Another remarkable case is Antarctica. This con
to drift further and further apart. But then, in the tinent has been glaciated for more than 20 million
early Cretaceous, the first angiosperms evolved and years, perhaps completely for 15 million years. But
spread to all continents eventually dominating most prior to that it was vegetated and in the Mesozoic is
of the ecosystems suitable for vascular plants. was a major centre for plant evolution, the pivotal
piece in the Gondwanan puzzle of a fragmented con
In the Late Carboniferous and Permian a group of tinent of which the pieces started to drift apart just
conifers known as Voltziales or Walchian conifers when modern conifers evolved. Palaeobotanists are
was widespread in Laurasia. Several had foliage discovering more and more fossil sites in Antarctica
leaves remarkably similar to some extant conifers, and other Gondwanan continents, yielding impor
but the ovuliferous cones and especially the seed- tant remains especially for the understanding of
bearing structures in these cones were very different. southern hemisphere taxa. A third phenomenon is
The famous studies of Rudolf Florin in the 1940s and the discovery of diverse conifer forests that existed at
1950s solved once and for all the century old debate very high Arctic latitudes such as Axel Heiberg Island
over the homology of the conifer cone and demon in Canada and Spitsbergen from the Cretaceous into
strated how the conifer cone has evolved, at least in the early Tertiary where now even tundra plants are
broad terms, from these Permian ancestors to, e.g., few and very small. In a warm temperate climate
present day pines. Modern conifers appear in the there existed mixed conifer forests reminiscent of
Triassic with Podocarpaceae and Araucariaceae, those occurring in, e.g., China today, but which had
of which some remains show traits that did hardly to adapt to four or five months without daylight in
or not at all change during all the ensuing mil winter. Many of these conifers were the deciduous
lions of years, for instance the structure of conifer ones we know today from far more southern lati
wood. Conifer wood is well preserved in numerous tudes, such as Metasequoia, Taxodium, Pseudolarix,
permineralized fossils allowing detailed study of cell and even Larix; the latter does occur in part within
walls. The wood of many conifers is anatomically the Arctic circle today.
relatively simple in structure and often hardly distin
guishable between families. The famous remains of Conifer phylogenetic systematics is marked by one
Late Triassic (ca. 210 million years ago) conifer tree predominant factor: extinction. Even the extant
trunks of the Petrified Forest in Arizona, USA could conifers reflect this very clearly, with 30 genera
have belonged to Araucariaceae, Podocarpaceae, (43%) represented by just a single species and a fur
Cupressaceae (Taxodiaceae) or more likely an extinct ther 11 genera with only two or three species, com
family. The big logs of these Triassic trees were trans prising 59% of the total of genera. Further analysis
ported by ancient rivers from a considerable distance of their relationships, as well as the fossil record, are
upland where they once grew to the place where they strong indicators that this lack of diversity at spe
now lay. Yet, these and other fossils demonstrate cies level is the result of extinction, not a reflection
abundantly how important and widespread conifers of incipient speciation. Take for example the Dawn
have been throughout the Mesozoic. Of particular Redwood (Metasequoia glyptostroboides), first found
interest is the conifer history on the Indian subcon as a Miocene fossil in Japan, then as a very restricted
tinent, which is now almost devoid of conifers (with living tree in China, and subsequently (described as
several species) from Late Cretaceous and Tertiary (Farjon, 2007). Indeed, cladistic analyses based on
deposits across the northern hemisphere even as morphology that sampled the fossil record did not
far north as the arctic islands of Canada. In 1994, give these results (Crane, 1985 and all subsequent
a conifer recognized as a third living genus in studies) and appear to be much more consistent
Araucariaceae, Wollemia nobilis, was discovered (Rothwell et al., 2009). There are perhaps analyti
some 100 km northwest of Sydney in a remote, deep cal problems inherent in the use of molecular data
canyon; it has almost certainly near relatives in the derived from organisms that only represent hugely
fossil record of Australia and beyond, but no more disparate lineages due to extinction. These difficul
than 100 mature trees survive miraculously in the ties will be only partly overcome with more compre
wild today. Similar stories abound in conifers, and hensive sampling of extant taxa (Mathews, 2009).
18 while the popular press hailed both examples as liv The inclusion of Gnetales in Coniferales (Pinales in
ing fossils when they were first discovered, that label Mabberley, 2008; Pinophyta of other authors) is,
would be appropriate to scores of other species. The therefore, premature at best and is not accepted in
fossil record of conifers is comparatively rich and this book. Nearly all recent molecular analyses of
very diverse, with numerous extinct species (Stewart the phylogenetic relationships among extant coni
& Rothwell, 1993; Anderson et al., 2007). Once fers divide them into two clades: the Pinaceae in one
important groups or families have become extinct clade and the remainder of the families in the other
long ago, others are at present merely represented (Cupressophyta). Araucariaceae and Podocarpaceae
by a few scattered species, but some, even though are placed in the latter clade as sister groups (e.g.
they evolved long ago in the Mesozoic, are still with Rai et al., 2008). In the fossil record, however, these
us and thriving. Although Florin (1951) considered two and not Pinaceae are oldest (Farjon, 2008) and
the Carboniferous Cordaitales to be ancestral to so we have again a conflict of evidence, albeit less
conifers based on demonstrated homologies, more serious than the gnepine controversy. Without con
recent analyses have cast doubts on this hypothesis sideration of the fossil record the relationships of the
(Rothwell & Serbet, 1994; Rothwell & Mapes, 2001). few remaining families of conifers are of only limited
Modern cladistic analyses based on molecular data academic interest.
(DNA) assessing extant gymnosperms usually con
clude on the monophyly of conifers (e.g. Stefanovi The 614 species of extant conifers are classified in
et al., 1998; Rai et al., 2008), but these studies often 8 families of which 541 belong to the three larg
suffer from undersampling of taxa. As a conse est families Pinaceae (231), Podocarpaceae (174)
quence of this, some extant taxa have been found to and Cupressaceae (135). Conifers are here consid
be closely related using molecular data (not available ered to include Cephalotaxaceae and Taxaceae; the
in fossils) while they appeared unrelated when mor Taxodiaceae are subsumed in the Cupressaceae with
phological data from the fossil record were included the exception of Sciadopitys (Japanese umbrella
with those from living taxa. The most controversial pine) which is assigned to its own monospecific fam
case of this lack of agreement in cladistic analyses of ily. Then there is the ancient family Araucariaceae,
diverse data sets involves the conifers and a group since the discovery of Wollemia with three gen
of gymnosperms known as Gnetales. The molecular era, and finally the still somewhat debatable family
evidence often indicates a close relationship of coni Phyllocladaceae (by some included in Podocarpaceae).
fers and Ephedra, Gnetum and Welwitschia, the three Within the species, subspecies and varieties are also
rather disparate and relict genera constituting the recognized, which brings the total to around 800 dif
extant order Gnetales. Despite seriously conflicting ferent conifers living today. Although such a figure
results among several of these studies (e.g. Rai et al., is not fixed, it gives a plausible estimate of the diver
2008; see for this conclusion also Mathews, 2009), sity at this level. As has already been hinted at above,
some analysts have boldly stated: Gnetophytes are behind this rather moderate number of taxa lies a
derived conifers and found them to be a sister group biological diversity far greater than in any other nat
to Pinaceae (e.g. Hajibabaei et al., 2006). Such con ural group of plants of comparable size. A long evo
clusions are superficial because they ignore the fossil lutionary history and ecological marginalization by
record and thereby crucial evolutionary information the generally more competitive angiosperms since
the middle Cretaceous have led to numerous adap Eurasian Nutcracker (Nucifraga caryocatactes) in
tations to cope with extreme environments, some Europe and P. albicaulis with Clarks Nutcracker
examples will be mentioned under the brief outlines (Nucifraga columbiana) in North America. Seeds
of families below. of these pines are wingless and will not fall out of
the cones but have to be pried out by a birds bill.
The Pinaceae, with 11 genera and 232 species are an Pine cones range enormously in size, from 2cm to
exclusively northern hemisphere family. No fossils 60cm in length and weights of a few grams to more
are known from the southern hemisphere or from than 2 kg in the amazing species P. maximartinezii of
parts that once were situated south of the equa Zacatecas, Mexico. This pine has a similar seed mor
tor. They are the ubiquitous pines, spruces, firs and phology as the known bird pines and the enormous
larches of northern boreal forests and mountain cones open their hard scales not enough to release 19
forests extending further south into cool to warm the seeds either. Which are the dispersing birds? We
temperate climatic zones. The pines (113 species) dont really know, but when I was once on the one
exhibit the greatest diversity within a genus of this mountain in the world where this pine occurs, I saw
family, both taxonomically and biologically. They some sinister ravens hopping about...could they
range from creeping shrubby species like Pinus have been the ones helping the tree regenerate?
pumila from NE Asian mountains with a cold mari Firs (Abies) are by comparison with pines much
time climate and P. culminicola on a few isolated more alike, but this book recognizes 46 species. They
mountain summits of NE Mexico to 60 or 80 meters are in many respects more demanding ecologically,
tall, straight pines like P. ayacahuite in Mexico and needing better soils and growing under climatic
P. lambertiana in California and Oregon. They occur conditions with less moisture stress and more equi
on the Arctic tree line of Canada and Russia (P. bank- table, cool temperatures. Hence, they have to be able
siana, P. sylvestris), with a winter of 8 months and to out-compete angiosperms, or be codominant in
-50 C temperatures, but also in steaming hot pine mixed forest. The seed cone bearing branches are
savannas on lowland coasts of the Caribbean and in situated in the top of the tree, where the beautiful
Thailand and Cambodia (P. caribaea, P. kesiya), where erect cones can freely receive windblown pollen and
tropical rainstorms in the wet season alternate with later disperse scales and winged seeds to the winds
frequent grass fires in the dry season. Growth rates that sweep through the crowns. Firs are mountain
can vary in pines as much as size. There are amaz trees, which are rarely extending to lowlands like in
ingly fast growers, such as P. palustris in the warm the northern taiga and there they are restricted to
SE of the United States, adding 23 m of height per more favorable sites for tree growth than other local
year, and pines that grow so slow that annual incre conifers. Spruces (Picea), with 38 species, are better
ment rings cannot be discerned without a magnify adapted to acidic soils and harsh climate than firs,
ing lens. Some of the latter, notably P. longaeva, carry although there are exceptions, such as Picea sitchen-
on for millennia. The oldest still living specimen sis which occurs within a narrow coastal strip with
tree in the world growing in the White Mountains much rainfall and moderate temperatures from
of California is more than 4806 years old (Lanner, Alaska to California. It reaches enormous sizes in
2007). It is the trees minimum age, an unknown the area around Vancouver and Seattle. Spruces can
number of years must be added for the young tree to form dense, uniform forests over very large areas,
have attained about 1.3 m height. It was a young vig a quality used to advantage in plantation forestry.
orous sapling of a few centuries when the pyramids Larches (Larix) with 11 species are deciduous, light
of Gizeh were being built during the 4th Dynasty of demanding pioneers that colonize areas after dis
ancient Egypt. While most pines disperse their seeds turbance, or occupy the most extreme habitats in
by dropping them from opening cones, to be carried peat bogs of the boreal zone or the Arctic tree line,
away by wind aided by wings, some have evolved a where on the Taymir Peninsula in Siberia at 75 N
strategy dependent on mutualism with birds which Larix gmelinii is the northernmost tree in the world.
carry the seeds to caches for winter storage, where Other genera have a limited number of species, e.g.
many of them germinate. There are even one pine- Pseudotsuga and Tsuga, or only one, like Pseudolarix,
one bird relationships, such as P. cembra with the a true relict now very rare in China (and not closely
related to Larix) and the enigmatic Cathaya, also in red or purple) receptacle to attract birds. The initial
China. cup-like seed scale grows around the seed to form
the epimatium, which in some genera (Afrocarpus,
The Podocarpaceae, by contrast, are a largely tropi Nageia, Prumnopitys) swells to a succulent fruit with
cal family, outside the tropics they occur mostly in the same attraction, taking over the function of the
mountains of the southern hemisphere. At pres receptacle. When a bird eats the fruit, the seed is
ent, 18 genera are recognized, with the largest, swallowed with the rest, but is protected from diges
Podocarpus, with 98 species nearly as speciose as tion by a hard seed coat and dropped somewhere
the pines (Pinus). Next is Dacrydium, with 22 spe away from the parent tree. In some species, like the
cies. The other genera all have fewer than 10 species. diminutive Microcachrys tetragona, a creeping dwarf
20 The family is undoubtedly of Gondwanan origin, shrub from the highlands of western Tasmania,
with fossils going back to the Triassic found on all most ovules develop after fertilization; the bracts of
southern land masses including Antarctica. In the the cone swell up and turn bright red so that a ripe
Podocarpaceae, taxonomic research is far from com seed cone resembles a little raspberry. Few studies
plete. On the one hand, more detailed observations are known that have dealt with the question of what
of morphology, now often backed-up by molecular animals act as dispersers; looking at these studies
evidence, has led to the recognition of more genera and what is known from pines, it is obvious that
to accommodate species formerly classified under in Podocarpaceae mutualisms are likely to abound
Podocarpus and to a lesser extend Dacrydium. On the and much is still to be discovered here. That is not
other, quite a number of species in Podocarpus are easy, because so many podocarps are scattered in
only known from a few herbarium collections, often tropical montane forests and there are so many spe
with insufficient material and little or no knowledge cies of bird and other fruit eaters around to be sus
of the biology of the plant. A comprehensive revision pected! As in Pinaceae, diversity of life forms is great
of this genus, and of the entire family, is wanting. A in this family, not only ranging from dwarfs like
critical taxonomic revision will most likely lead to a Microcachrys to giants like Podocarpus totara of New
reduction of the number of species recognized, but Zealand, but even including a true parasite, the only
there may also be new discoveries in under-collected one in gymnosperms. Parasitaxus usta, from New
regions, such as western New Guinea and southern Caledonia, lives on the root bases of another podo
Venezuela. For this book, only a preliminary revi carp, Falcatifolium taxoides. Some species are shrubs
sion of the family was possible, so some relatively well adapted to nutrient-poor and harsh environ
recently described species have here been retained ments, such as the 3 species of Lepidothamnus, one
with the benefit of the doubt. The Podocarpaceae are of which grows in peat bogs in southernmost South
characterized by specialized seed cones, in which in America, the two others in the mountains of New
most cases during ontogenesis the number of seeds Zealand. There is also a true rheophyte, the dwarf
is reduced to one, which becomes variously enclosed tree Retrophyllum minus from New Caledonia, with
or subtended with soft, often colourful and at any thick spongy bark, perhaps to protect it from rocks
rate to certain animals tasteful tissue. As a result, and other debris smashing against it in flash floods.
these conifers are adapted to dispersal of seeds by Others are emergent trees in tropical montane
animals (zoochory). However, as in other conifers, rain forests, such as many species of Podocarpus,
the Podocarpaceae are wind pollinated. Hence, the or shrubs to dominant trees in the moss-forests of
pollen cones of the Podocarpaceae do not differ from high tropical mountains, like species of Dacrydium
types found in the Pinaceae, but their seed cones are in New Guinea.
profoundly different. With few exceptions, a com
pound cone structure, with an axis beset with bracts The Cupressaceae are the only cosmopolitan family
and axillary ovuliferous scales, can only be discerned of conifers at present. The family is currently recog
in the initial (pre-fertilization) phase. As soon as the nized to have 30 genera and 135 species. This count
first ovule, e.g. in Podocarpus, has been fertilized, the includes the former, well-known family Taxodiaceae,
other ovules are aborted and the bracts swell and a family concept which cannot be maintained when all
fuse together to form a juicy and colourful (often the information now available, including phylogeny,
is considered. The species accomodated in the components of subalpine scrub communities, and
Taxodiaceae were in fact a loose assembly of relict the diversity in this family, and that of the conifers in
taxa having retained a few primitive traits, such general, will become clear.
as spiral arrangement of leaves Metasequoia, with
decussate leaf arrangement (phyllotaxis) is by many The smaller families, except the Taxaceae, have a
other characters related to Glyptostrobus, Sequoia more limited distribution. The Araucariaceae occur
and Taxodium with spiral phyllotaxis. Decussate and in South America, with two species, one of which is
whorled phyllotaxis, as well as other leaf charac the well known Monkey puzzle Araucaria araucana,
ters, appear to have evolved more than once from but is more abundant in Borneo, New Guinea, NE
a presumably ancestral spiral arrangement. While Australia, and New Caledonia and other SW Pacific
in the Podocarpaceae extreme reduction of parts has islands. The genus Araucaria was in the Mesozoic 21
led to a single-seeded cone, in Cupressaceae reduc nearly cosmopolitan, and hence the typical coni
tion of another kind has taken place. Here, the seed fer with which to depict the dinosaurs, but became
scale itself has been lost or, in some cases, e.g. in increasingly restricted in its distribution. It does not
Cryptomeria japonica, vestiges in the form of seem mean that all the 19 species currently recognized are
ingly functionless appendages of what may have living fossils; it is more likely that most of the spe
been a scale, remain. The erect ovules (most other cies in New Caledonia, where at present the genus
conifers have inverted ovules) originate on the apex has its centre of diversity with 13 species, evolved
of a shoot in the axils of transformed leaves (bracts). there more recently. Yet others, such as A. araucana
These bracts enlarge in various ways and ultimately and A. angustifolia from South America and A. bid-
form the cone scales, covering the seeds. willii from Queensland, Australia, may have changed
The worldwide distribution of Cupressaceae is little since the Jurassic. In contrast with Araucaria,
reflected in the great ecological diversity among Agathis, with 17 species, is restricted to Malesia, NE
its species, although only a few are tropical such as Australia and islands of the SW Pacific. It is not yet
some species of Callitris and Libocedrus from north known in the fossil record prior to the Tertiary, nor
ern Australia and New Caledonia and Papuacedrus from other land masses. The most famous mem
papuana from New Guinea and the Moluccas. The ber of this genus is the Kauri, Agathis australis, of
other species occupy all the major environments the northern part of North Island, New Zealand.
that conifers thrive in generally, but in addition have The third genus, Wollemia, was discovered in 1994
many species well adapted to semi-arid conditions. in a hidden canyon in Wollemi National Park, New
Especially diverse in semi-arid environments, which South Wales, Australia, and is one of the most inter
occupy large parts of the land masses of four conti esting of recent conifer discoveries. It is considered
nents, are Cupressus and Juniperus in the northern to be another living fossil and true fossils are indeed
hemisphere and Callitris in Australia; some of the now being assigned to this new genus.
species even occur in true deserts. Other extremes
are altitude records, e.g. Juniperus indica found at The Phyllocladaceae are distinguished by greatly
5100 m altitude on the moraines of Tibetan glaciers reduced true leaves and phylloclades, representing
coming from Mt. Everest. Other Cupressaceae thrive planated shoots, taking on the tasks of assimilating
in ecosystems that can hardly be more different leaves. They also have a distinct system of pollen cap
than these: the wet, oceanic forests of giant conifers ture by the ovule, which sets them apart from most,
along the Pacific coast of North America from cen but not all podocarps, as does the formation of a true
tral California to Alaska. Sequoia sempervirens from aril around the seed, as in the Taxaceae. However,
California is the tallest tree in the world, reaching molecular evidence appears to indicate that they
up to 117 m. Sequoiadendron giganteum, confined to are very closely related to the Podocarpaceae. The
the western slopes of the Californian Sierra Nevada Phyllocladaceae are from a Gondwanan ancestry
a little further inland, is a living fossil of gargantuan and greatly reduced. The four species of its single
proportions. Compare this with the dwarfed spe genus are now scattered in Malesia, New Zealand
cies Microbiota decussata from the Russian Far East and Tasmania. The Sciadopityaceae are represented
and Diselma archeri from Tasmania, both important by a single species, Sciadopitys verticillata from a few
localities in Japan. This conifer has until recently genus Taxus, here recognized with 10 species, but
been misunderstood; it is not a pine and although recently again the subject of debate, when a long-time
related to the Cupressaceae it has a whole suite of student of the genus raised the number of species to
unique characters. It is a prime example of a relict 24, with 55 varieties (Spjut, 2007). In the Taxaceae, 4
taxon and deserves more intensive study by all disci other genera are recognized, of which Amentotaxus
plines relevant to its taxonomy. The Cephalotaxaceae, and Torreya have 6 species. All are evergreen shrubs
with a single genus as recognized by most taxono or small trees adapted to live in the understorey of
mists, accommodates 8 species, all eastern Asian, tall forests and have developed a single arillous seed
ranging from Korea and Japan to Malaysia. In some of which the aril is either partly covering the seed and
DNA-based analyses, this family has been proposed soft and juicy and colorful red or yellow, or wholly
22 for synonymy with Taxaceae, but I think there are covering it and more fleshy and bluish green to pur
arguments to retain it as a separate family. Several ple. The aril is eaten by birds which disperse the seeds
species are cultivated widely and the taxonomy is still undamaged. In recent years, the discovery of chemi
somewhat unsettled, in part because some species are cal compounds (taxanes), which have proven to be in
based on cultivated plants, in part because species certain cases effective against cancer in humans, have
have been based on variable foliage characters due to focused a lot of research on the species of Taxus. This
the extremely reduced ovuliferous cones. A similar event has also demonstrated how much we can still
difficulty besets the Taxaceae, and especially its largest learn about conifers.
T H E D I S T R I BU T IO N A N D E C O L O G Y O F C O N I F E R S
The 614 species of conifers cover a large proportion while the Greenland icecap prevents any form of
of the land surface of the earth. The map on p. xxx vegetation. The steppes of North America, Central
shows that the greatest covering of land by conifers is Asia, Tibet and Mongolia, parts of Africa, and
clearly to be seen in the northern hemisphere. This is Patagonia in South America can have some conifers
due to the extensive conifer forest of the boreal zone here and there, but are also largely without them. 23
in Eurasia and North America. There are only few Although a few conifers occur in deserts, most of the
species occurring there and the diversity increases worlds deserts, i.e. the deserts of central and western
dramatically further south, while the areas being Asia, Arabia, North Africa (Sahara), and Australia,
covered decrease. The area indicated to be covered are devoid of conifers. Another vegetation type
by conifers on the map is, of course, not uniformly mostly devoid of conifers is lowland tropical rain-
covered with conifers, they usually occur together forest, such as the Amazon Basin in South America
with angiosperms or grow in patches too small to and the Congo Basin in Africa but also in smaller
separate on this scale. There are already big gaps at areas like the Yucatn Peninsula in Mexico and the
around 30 North, which continue across the equator southern lowlands of New Guinea. Other large areas
into the southern hemisphere. In the far south there without conifers have been stripped of their natural
is much less land, but except Antarctica which has vegetation and are now occupied by agriculture and
been omitted from the map for obvious reasons it urbanization, especially in Western Europe, India,
is quite well covered with conifers, with few big gaps. Bangladesh and northern China. However, some
Conifers are present in nearly all the major veg- of the remaining areas devoid of conifers cannot be
etation types of the world. They are absent from only resolved with an ecological explanation. Large parts
a few and are very rare in some others. In the far of South America and larger parts of Africa and the
north, a few conifers occur in tundra vegetation, Indian subcontinent are the major gaps in conifer
mostly in the transition zone from boreal forest to distribution that have no explanation in terms of
tundra. In the High Arctic of northern Canada and unsuitable climate or soils. Did these parts ever have
northern Siberia (cold deserts) they are left behind, conifers, and if so, why did they loose them?
The global distribution of conifers shown on a world map with approximate equal area projection. The
black areas on the map are not uniformly covered with conifers; they occur together with angiosperms or in
patches too small to separate on this scale.
Discounting the Mediterranean coastal areas, which origin. Such widely distant occurrences of taxa in
biogeographically belong to Eurasia, Africa is the the Cupressaceae seem to throw the separation of
poorest continent for conifers today. In Sub-Saharan extant conifers into northern and southern origins
Africa we only have Widdringtonia (Cupressaceae) into doubt. Araucaria may have originated before
with four species, Afrocarpus (Podocarpaceae) with the separation of Pangea into two super-continents,
five species and Podocarpus with four species. Vast but as far as we know, the Cupressaceae evolved after
areas of Sub-Saharan Africa have no conifers at all. that event.
The Indian subcontinent south of the Himalaya and
adjacent hills has only a single indigenous conifer: One final, intriguing observation about present-day
Nageia wallichiana (Podocarpaceae). There are plenty conifer distribution is that more than half (ca. 330)
24 of suitable areas for conifers in both, as the successful of all species occur on the Pacific Rim. All families
plantation of conifers for forestry in Africa and India and 83% of all 70 genera are represented. The Pacific
demonstrates. The causes are most likely of a histori- Rim is the system of mountain ranges and islands
cal kind, having had effect over time spans on a geo- around the Pacific Ocean in both hemispheres,
logical scale. It appears that isolation, caused by the mostly forming part of so-called subduction zones
break-up of the southern supercontinent Gondwana where oceanic crust plates slide beneath the conti-
and the increasing separation of its constituent land nents, causing volcanism and thrusting up of moun-
masses, is a major factor behind the gaps in conifer tains and islands in the process. Going clock-wise an
distribution of land masses of Gondwanan origin. starting at 10 oclock the diversity centres for conifers
Connections between land masses remained much are Japan, the Pacific North-west of the United States
longer in place with the break-up of Laurasia, the and Vancouver Island, California, southern Mexico
northern super-continent, which also became less and Guatemala, southern Chile, New Zealand,
fragmented. Both dispersal and vicariance played a Tasmania, New Caledonia, Fiji, the coast of New
part in the history of conifer distribution leading to South Wales and Queensland, New Guinea, Borneo,
the present situation. However, due to the size, prox- the Philippines, and Taiwan. All of these areas have
imity, and orientation of land masses of Laurasian many species in several genera and families. In con-
origin, all situated in the northern hemisphere, dis- trast the coastal areas of the Atlantic and Indian
persal accounts for much of the distribution of gen- Oceans do not have such diversity of conifers, with
era and species. Losses in the past could often still be the exception of Morocco, which has a moderate
made up for by new arrivals. diversity of 12 species, and Florida, with 10. The
The long history of the assemblage and subse- explanation is probably again historical: none of the
quent fragmentation of the two super-continents, other oceans are nearly as old as the Pacific Ocean,
Laurasia in the north and Gondwana in the south, is which has therefore provided coastal mountains and
apparently still reflected in the distribution patterns islands suitable for well moisturized montane forests
of conifers across the globe. If we look at the dif- from well before the origin of the angiosperms.
ferent taxa this is even more clearly demonstrated.
At the family level, Cephalotaxaceae, Pinaceae, In the north, around the Arctic Circle, the conifer
Sciadopityaceae and Taxaceae are Laurasian, whereas forest or taiga begins where the tundra ends. The
Araucariaceae, Phyllocladaceae and Podocarpaceae taiga is not entirely homogeneous. It is interrupted
are Gondwanan, although Araucaria once extended by lakes and swamps and dissected by large riv-
into Laurasia. Cupressaceae as a family are cosmopol- ers. There are few conifer species and most of them
itan, but the 30 genera that make up this family today belong to the family Pinaceae. In some areas two or
are divided in northern and southern hemisphere three of these conifer species may occur together, but
groups, with a few trespassers, some only known very often the forest is formed by only a single spe-
from the fossil record. Some of these exceptions cies. The environment of these conifers is dynamic.
are intriguing enough, though. The South African Disturbance by fire and flooding is an integral part of
genus Widdringtonia has been found in the Upper the taiga ecosystem, and natural disturbance, mostly
Cretaceous of North America and Austrosequoia in the form of fires, often covers large areas. Storms,
wintonensis from the Cretaceous of Australia is very however, are rare. The air burst of a large meteoroid
similar to Sequoia or Sequoiadendron of Laurasian or comet fragment in June 1908 in Siberia, known as
the Tunguska Event, destroyed large tracts of taiga conifers have a natural chance. Once they are mature,
forest. Outbursts of defoliating insect plagues, often they will persist because they now can compete with
associated with freak weather conditions, can lay any angiosperm. One of the key factors for the suc-
waste to the forest as well. However, usually within a cess of conifers on poor soils must be the extensive
few decades the conifer forest is coming back from development of mycorrhizal symbiosis with cer-
abundant seed. It was their capacity to conquer new tain fungi, which is especially prevalent in conifers.
land in great numbers that enabled conifers to return This form of collaboration between entirely differ-
and to occupy the vast northern regions during the ent organisms enables conifers to cope with very
relatively short interglacial phases between the ice low levels of essential minerals such as phosphorus
ages of the Pleistocene. But the present conditions and nitrogen in the soil, because the fungal hyphae
are still harsh, the growing season is very short, and greatly expand the surface and thereby uptake of 25
only a few species have been able to adapt to these. water and nutrients by the plant root system. It is
now even being thought that plants would not have
Conifers predominantly occur on soils or in climates succeeded to come onto dry land permanently
that are sub-optimal for plant growth. However, as without the assistance of these fungi; conifers were
we know well from plantation forestry and horti- among the first to succeed in leaving the lakes and
culture, there is no evidence that conifers prefer swamps behind them.
nutrient-poor soils or harsh climates. Many species
that are restricted to these poor conditions in their Although it is the most widely applied strategy, not
natural habitat grow exceedingly well if planted on all conifers are escapists. Especially in areas on the
fertile soils. Monterey or Radiata pine (Pinus radiata) Pacific Rim, there are many conifers that grow very
has been planted on a large scale especially in the tall, rising well above the general forest canopy. Most
southern hemisphere and is one of the most produc- of them can live very long, some even thousands of
tive trees in the industry. Yet its natural habitat is years. These giant trees appear to be scattered in the
nutrient-deficient podzolic sand in the salty winds forest as veteran trees, or they occur in cohorts of
of the Pacific Ocean right on the Californian coast. more or less even-aged trees when still of a lesser
Many species of conifer are more or less restricted to age. These forests grow on the whole on mineral-
ultramafic soils derived from serpentine or similar rich, young soils, often of volcanic and sometimes
rocks, where phosphorus is almost absent. It appears of glacial origin. Competition among abundant trees
that conifers, in order to survive in nature, have of many species is fierce. The ecological strategy here
often adapted to such poor sites. It is the strategy is closely tied in with localized forest disturbance.
of evasion. What are these conifers trying to escape Fires, land slides and destructive storms in these
from that they have adapted to put up with such places occur infrequently, sometimes with intervals
poor conditions? The answer is competition from of several centuries. The old, big trees are the survi-
angiosperms, in particular fast growing, large-leaved vors of such disturbances. The bigger the tree, the
herbs, shrubs and trees. While conifers, once they more resistant it becomes; the bark of huge trees of
are full-grown trees, can often outgrow angiosperm Sequoiadendron giganteum from the western slopes
trees, they often have a slower start. It has long been of the Sierra Nevada in California is up to 60cm
assumed that the evolution leading to weedy herba- thick and a very effective protection against forest
ceous angiosperms has caused the decline of coni- fires. Nearly all of these giants among the conifers
fers. Grasses are particularly effective in suppressing are light demanding and would not successfully
juvenile conifers and have excluded conifers from regenerate under the canopy of other trees, being
the steppes in Asia and the Americas and probably either angiosperms or conifers. These trees have to
from the grasslands of Africa. That conifers can be removed first. On open sites, regeneration of the
grow well there is again demonstrated by plantation big tree conifers is often abundant, but soon compe-
forestry. Overgrazing and fire prevention, suppress- tition starts and a struggle for space, light and water
ing the grasses, can lead to the return of conifers; begins. Since some of the big conifers can grow
this happens at present with junipers (Juniperus) taller, they have a chance to become dominant above
on the rangelands of the American West. However, the canopy of the other trees. The giant ultimate sur-
if site conditions are poor for any plant growth, the vivors are the insurance policy of the species.
There are many mutually beneficial relationships of seed dispersal. Probably all 174 extant species in
between animals and plants, almost all of which have the family have their seeds dispersed by birds. In the
developed with angiosperms on the plants side of the Cupressaceae, it is the genus Juniperus in which the
deal. Pollination provides numerous examples and it cone, in a very different way, has become fruit-like.
is the single reason why flowering plants developed Birds are again the main dispersers of the seeds, in
their almost endless variation of flower designs. Not many species reduced to one large seed per cone;
a single conifer has ever managed this as far as we quite a reduction in number if the evidence is cor-
know, they are all wind pollinated, which is, lack- rect that the genus may have arisen from an ancestral
ing precision in pollen delivery, a wasteful process. species of Cupressus, of which many species have well
Wind pollination has, however, not prevented them over 100 seeds per cone. The seeds of all species in
26 from becoming abundant and widespread, and the Taxaceae arise at the apex of dwarf shoots, a seed
some wind pollinated angiosperms, like grasses, cone does not even develop to become aborted in a
have done well, too. Many conifer species have been later stage as in some species of the Podocarpaceae.
more successful with seed dispersal aided by ani- An aril, arising from the initial seed integument,
mals. In general, there are two scenarios for seed takes on the fruit function. The imitation of angio-
dispersal: (i) the animal transports and eats a fruit sperm fruits has proved very successful in dispers-
containing seeds, which pass through the animal ing the seeds of conifers with a minimal waste of
undigested, and (ii) the seeds are transported and resources. In the second scenario of seed dispersal
hidden to be eaten later and not all hidden seeds are by animals, some species of pine (Pinus) have devel-
eaten. Conifers have adapted to both scenarios and oped mutualistic relationships with birds of the crow
the first one is more common than the second. The family (Corvidae) to effect seed dispersal. The birds
families Podocarpaceae, Taxaceae, Phyllocladaceae, remove the seeds from the specially adapted cones
Cephalotaxaceae and Cupressaceae all have many and hide them in the surrounding terrain for later
species in which the seed cone, or the seed, has consumption. Large quantities of seeds are involved
evolved to resemble a fruit. In all cases, the cone and invariably not all seeds get eaten. Where their
has been extremely reduced, in some cases it effec- close relatives have winged seeds, the seeds of these
tively disappeared. With few exceptions, the num- pines are wingless; wings would be useless and a
ber of seeds has been reduced to one only. In many waste to produce. In this way, the pines can travel
ways, the Podocarpaceae, the second largest family upslope independent of prevailing wind conditions,
of conifers, has been the most innovative as well which for some of the subalpine species may turn
as the most successful family to adopt this strategy out to be an advantage during a warming climate.
T H E E C O N OM IC I M P O RTA N C E O F C O N I F E R S
The economic uses of conifers can be conveniently subsect. Strobi), are providing wood of higher qual-
divided in those centring on wood, other derivative ity with properties such as high dimensional stabil-
products, and horticulture. The first two are mostly ity, straight grain, softness and workability together
industrial. Horticulture does not generally manu- with large, straight dimensions especially when
facture products but focuses on the cultivation and taken from old growth forests. These timbers are 27
trade of the whole plant and has a strong element of applied in the building industry for, e.g. doors and
non-commercial interest. However, there are mani- windows, in furniture making and even for musi-
festations of both applications evident in all three cal instruments such as organ pipes and piano keys.
categories of economic use of conifers, so that the Firs (Abies) produce lightweight, relatively soft,
demarcation between industry and leisure is not an creamy white to pale brown wood; the high grade
absolute one. It could even be argued that these dis- timber from forests with these species is sawn for
tinctions have become less clear in recent times than framing material and for plywood and veneer. The
they were in the past, and examples will be given in wood of spruces (Picea) differs markedly from that
this chapter. of firs and is consequently used for different, mostly
Today the world trade in coniferous wood is huge. less refined purposes. In northern lands, it was and
Conifers provide about 60% of all wood used for often still is the principal tree for the construction of
industrial purposes. They dominate industrial wood log houses. In Norway, the ancient village churches
supply because of both technical and economic were entirely made of wood, often a mixture of
advantages over wood from angiosperms. The vast spruce (Picea abies) and pine (Pinus sylvestris); some
reserves of natural conifer forest in the boreal zone date from nearly 1000 years ago and are still intact.
of the northern hemisphere play an important part In the Alps similar uses in construction are made
in this supply, but plantations, especially in the of larch (Larix decidua), as its wood is particularly
southern hemisphere, of particularly pines take up resistant to weather and rot. All three genera also
an increasing proportion of this. Nearly all this mass provide pulpwood for the paper industry; the trans-
production goes into pulpwood for the paper indus- parent windows in envelopes are a paper made from
try, although about of conifers in plantations in the larch wood. In some genera of conifers, various cir-
world are destined for timber. The more specialized, cumstances have caused only one of the species to
high quality woods used for all the versatile applica- become of high economic importance. An example
tions mentioned in this chapter come from forests is the Douglas fir (Pseudotsuga menziesii), which has
that can be allowed to grow older. In the trade, the become the most important timber tree in North
wood of conifers is known as softwood and that of America. It has consequently been introduced by
angiosperms as hardwood. These terms are mislead- foresters in many temperate regions around the
ing if applied to all conifers, because many species world. Douglas fir grows rapidly, is straight and tall
produce very hard wood indeed, while some of the and produces large volumes of wood per hectare,
softest woods in use do not come from conifers but especially in managed forests where competing spe-
from angiosperms. By far the greatest volume of cies are excluded or suppressed. Its wood is used for
industrial wood is produced by species in the fam- plywood and construction, both exterior and inte-
ily Pinaceae, which occur naturally in the northern rior, and it has a reasonable durability for outdoor
hemisphere only, but have been widely planted for applications such as telephone poles and railway
this purpose in the southern hemisphere. In par- sleepers. The wood of the Cupressaceae differs mark-
ticular, the genus Pinus stands out, with several edly from that of Pinaceae. It is more fibrous and
species being among the fastest growing plantation contains less resin; it is also mostly very decay-resis-
trees producing pulpwood on short-term rotations. tant and many species have fragrant properties due
Several other species, especially white pines (Pinus to volatile chemical compounds. These properties
make it highly desirable in China and Japan, where family are often selectively logged for timber. The
it was traditionally used in the construction of tem- most important genus, Podocarpus, is also the most
ples and other ceremonial buildings. Some species widespread; of regional importance are Afrocarpus
have been over-exploited and good sized trees are (Southern Africa), Dacrydium and Dacrycarpus
now rare. Other species, like Cryptomeria japonica, (Australasia) and Nageia (Southeast Asia). All yield
have been widely planted, in plantations in Japan, light to medium-weight, pale coloured wood, known
where it is native, as well as in China and Taiwan. as podo in the trade, with a straight grain and even
Some of this cupressaceceous wood splits easily into texture that is easy to saw and plane but is often brit-
shingles, and its rot-resistance was noted by people tle. It is not durable when exposed to the weather, so
in regions as far apart as the Pacific coasts of Canada its building applications are for indoor construction
28 and the northwestern United States, Chile, Japan and only. High grade timber can used for door and win-
Viet Nam, where these shingles were traditionally dow frames, panelling, veneer, cupboards, furniture,
used to cover the roofs of houses. Durability has cabinet work, joinery, household utensils and engi-
also been the major property that made juniper neering instruments like drawing boards and rulers.
(Juniperus) the tree of preference for fence posts, but The New Zealand species Podocarpus totata provides
metal is pushing it out of the market. Some species the only softwood that is resistant to attack by marine
in the Cupressaceae produce wood with beautiful borers, so it was used for ship and boat building as
patterns and are therefore prized for the making of well as wharf building and harbour construction until
cabinets and other pieces of furniture. In particular, a ban on further logging stopped it. The Maori built
the large, ancient coppice stools of the North African their famously long war canoes with the wood of this
species Tetraclinis articulata are valuable and were large indigenous tree.
already sought after by the Romans. Few conifers
have this coppicing capacity, i.e. re-growth from a Of second industrial importance to wood of conifers
stem base after repeated cutting. Another source for is their resin. Resin is present in all conifers, albeit
this type of wood, suitable also for wood turning, is not in equal quantities. Resin in leaves can be dis-
yew (Taxus), which is hard, dense, heavy and resis- tilled from them, resin in wood can be tapped as well
tant to decay. Perhaps most famous were the English as distilled, and there is even resin to be mined. The
yew longbows of the Middle Ages; the arrows shot resins of conifers are mostly terpenoid, with some
from these bows could penetrate a knights armour phenolic resins (Langenheim, 2003). Only two fami-
at 200 meters and helped the English win the battles lies, Araucariaceae and Pinaceae, produce copious
of Crcy, Poitiers and Agincourt in the Hundred amounts in the wood, where the resin is stored in resin
Years War (13371453). ducts or canals. The genus Agathis (Araucariaceae)
The most tropical of the conifer genera, Agathis is the most copious producer of resin; over centu-
(Araucariaceae) is one of the most valued timber ries, resin has flowed from the trunks of large trees
trees in Australasia. The wood known as kauri in the onto the ground where it has accumulated, forming
timber trade is light and soft, pale yellow or straw- large deposits of copal which can be excavated. Its
coloured, often with darker heartwood ranging use is mostly for varnishes. More commonly, resin
from pink to dark red brown. The wood of Agathis is tapped from the trunks of pines. Major resin pro-
has many uses, from indoor construction, panelling, ducing species are, or were, Pinus kesiya in Southeast
boat masts, joinery, furniture, pencils, matches and Asia, P. massoniana in China, P. pinaster in Europe,
matchboxes, rulers, and piano parts. Naturally it is especially France, and P. palustris and P. elliottii in
excellent for plywood and veneer, while more indus- the Southeastern United States. The resin tapped
trial uses of lower grade wood are pulp for paper forms the basis of many products in industry, such
manufacturing and high grade charcoal. In Indonesia as turpentine, rosin and pitch together known as
and Malaysia Agathis is exploited heavily for export naval stores oils, varnishes, printing inks, sealing
of raw timber (round logs); in the Philippines this wax, soap, plastics, and fireworks. Coarser prod-
has already led to a total ban on further cutting, ucts are obtained by destructive distillation of res-
while export is banned from Papua New Guinea. In inous wood. In the age of wooden ships, pitch and
tropical countries the wood of the Podocarpaceae tar were indispensable to keep them seaworthy by
is usually highly valued and trees belonging to this caulking the seams with pitch and by tarring the
rigging. The term naval stores for these and similar rosin as incense was a privilege of priests and kings.
products dates back to the 17th century, when the Religious or symbolic use also lies at the root of the
English navy required large quantities of these for an Christmas tradition to decorate a room in the house
expanding fleet. with an evergreen tree and the growing of several
species of conifer for this purpose has developed in
The value of conifers for human food is marginal and the last 150 years or so to a significant industry.
consists almost exclusively of the seeds of several
species of pine (Pinus) and three species of Araucaria, Conifers in horticulture certainly started as a pas-
two in South America and one in Australia. Pine time, not as a business. Its earliest roots lie undoubt-
seeds are nutritious and sometimes tasteful food edly in China and Japan, where planting of trees for
and some are of commercial value today. In Europe, aesthetic reasons antedates that activity in Europe by 29
it is mainly the Mediterranean species P. pinea or centuries. In that tradition, form, shape and colour
umbrella pine which yields seeds of good size and were of greatest importance as part of a reconstruc-
taste for the food market (Italian pignolia), used in tion in situ of idealized pictorial land- and town-
processed food as well as sold as whole seeds. The scapes. By contrast, in Europe the planting of (exotic)
distantly related Asian pine P. koraiensis produces conifers in gardens and parks mostly resulted from
similar kernels and has become the leading species a curiosity about the natural world of distant lands
in the export market. Pinus gerardiana, or chilgoza, and the products these brought forth. This devel-
and P. bungeana, or lace-bark pine, are other Asian oped in the 19th century to a rage among the landed
pines with good edible seeds. In the United States gentry of Europe to obtain species, which led them
and Mexico there are several species with edible to employ plant hunters to travel to America and
seeds, of which P. cembroides, P. edulis and P. mono- Asia to obtain suitable tree seeds that could be
phylla are the most widespread, which are com- grown in Europe. Horticulture developed in part
monly known as pion or pinyon pines. In Russia, from the demand that this trade generated, with
P. sibirica produces seeds which are harvested for oil nurseries in e.g. England and Germany specializing
extraction. The seeds of Araucaria, although deli- in trees, including conifers, for large gardens, parks
cious when roasted, are more of traditional value to and estates. This again fed into plantation forestry,
indigenous people than commercialized for the food when some land owners saw potentials for planting
or delicatessen markets. The rarity of the trees of A. timber trees on their estates. By the middle of the
araucana and A. bidwillii due t o past over-exploita- 19th century, conifers from North America were well
tion prevents large scale seed harvesting. Flavouring established and common, while the first introduc-
is probably best known from the use of juniper tions from China and Japan had just arrived. About
berries (seed cones) in producing gin. The Dutch 6070 years later, these had become common, too. It
words for gin and for the shrub Juniperus communis was this widespread planting of exotic conifers and
are, respectively, jenever and jeneverbes, in the lat- other trees which caused a demand for descriptive
ter the Dutch word for berry is added. This species, literature and gave rise to a number of manuals or
together with its varieties, is the most widespread handbooks, particularly in England and Germany.
conifer in the world and the various brands of gin, Naturally, these books concentrated on the species,
like aquavit, gin, jenever and Schnaps, are all made and soon also cultivars, that were grown in Europe.
with it. The resin of the Mediterranean pine Pinus The British Isles, due to a relatively mild but varied
halepensis is used to flavour retsina, a Greek white climate and varied geology and topography, as well
or ros wine popular with locals as well as tourists. as widespread private landownership that has con-
Scent is close to flavour. The typical fragrance of a tinued to the present, are the worlds centre of diver-
northern conifer forest is due to the volatile com- sity for introduced conifers. Current membership
ponents of the terpenoid resins of the conifers and of the International Dendrology Society (which was
these are being used in the cosmetics industry. The established on the continent) reflects this. However,
volatile components were and are also used in reli- in recent decades other parts of the world have been
gious ceremonies, as fragrance and scent play a catching up, such as the United States, Australia and
major role in imagination. The Aztecs called teocote New Zealand. With parts of these countries in a
pine (Pinus teocote) the pine of the gods; to burn its warmer and either wetter or drier climate than most
of Europe, the range of species that can be grown disillusion may have been that dwarf conifers often
extends beyond the more or less common assortment grow bigger after a slow start than they were prom-
used in Europe. There is no good reason why tropi- ised to do when planted. The methods and practices
cal conifers should not be planted in gardens and of growing dwarfs only slowly developed into what
parks of tropical countries, which is one reason why professional nurserymen can achieve in this mtier
this book includes all species. While the planting of today. The trade is, of course, influenced by the
conifers in gardens and private parks remains pas- demand but also creates it by making new and bet-
time activity, horticulture with conifers has devel- ter cultivars available. Many good dwarf and colour
oped into a substantial business aiming at a different cultivars have been developed in the last decade or
and wider market. In recent years the trade appears so. A major development has been the utilization of
30 to have been emphasising dwarf conifers, but the witches brooms occurring as a result of DNA muta-
fashion is not new. One of the many garden crazes in tions in buds or shoots. Unlike those resulting from a
the Victorian age was building rock gardens in parks trees reaction to parasites or pathogens, these genetic
with plantings in it that ought to grow slow and mutations alter growth of any shoot taken from the
remain moderate in size. In Britain, dwarf conifers witches broom, resulting in clones with identical,
became popular for a time from the 1850s onward. permanent dwarf growth characteristics. Although
Interest was renewed in the 1960s but this time it they were first used as long ago as 1836, in recent
was a fashion to plant heathers in suburban gardens times selections derived from witches brooms have
and it affected Europe as well as the United States. become a major source of dwarf conifers in cultiva-
Consequently, many arboreta and pineta followed tion, and a constant stream of new cultivars is enter-
suit and established heather gardens with small or ing the market. With a growing knowledge of causes
medium size, preferably columnar or fastigiate coni- and processes, especially genetics, and improved
fers. When people began to be bored with this, the techniques in propagation and cultivation, we can
heather garden went out of fashion again, although see this develop in future into an industry where
it may still have its adherents. One reason for the conifer cultivars are being made as well as grown.
T H E C O N SE RVAT IO N O F C O N I F E R D I V E R SI T Y
Extinction is an apparent fate of all species. What Against this rather overwhelming perspective, it
concerns conservationists is the current high rate of could appear to be next to hopeless to take action
extinction, well above what is believed to be a back- for the conservation of conifer diversity in the natu-
ground rate. Different estimates of these rates exist ral habitats in which they occur. The real big issue
and are debated, but what is undisputed is that the is human demography, but to address that is both 31
current rate is abnormally high and that humans complicated and difficult. It has to be addressed,
are causing it. The primary cause of the decay of or else the size of the population will be corrected
organic diversity is not direct human exploitation naturally, which will be disastrous. Meanwhile, we
or malevolence, but the habitat destruction that can and should buy time, although there is not much
inevitably results from the expansion of human time left. This is why the conservation of species in
populations and human activities. This is how Paul their natural habitats is very important, and even
Ehrlich, a well known American ecologist and writer more urgent when certain species are keystone in
on the biodiversity crisis, has summed up the cause the ecosystem. Many conifers perform that role,
of the present mass extinction of species (Ehrlich & because they are large, long living trees providing
Ehrlich, 1981). He called the destruction inevitable habitat for numerous other species. Conservation
and that is what it is under those circumstances. He of these conifer species in functioning ecosystems
later calculated that by the end of the 1980s humans in the wild means the conservation of many other
had consumed, directly and indirectly, up to 40% of species. It is therefore a good strategy to concentrate
the total net primary production of stored biologi- conservation efforts on conifers. Especially in the
cal energy produced by plants from solar energy. tropics, however, our knowledge of the role of coni-
So, one species is using two fifths of all biological fers in ecosystems is rudimentary. Where natural,
production on the planet. He warned that when the undisturbed forests are formed by many tree species,
human population doubles again, that could become conifers often occupy restricted localities and occur
as much as 80%. Logically, if human activities and there in numbers. The ecosystem will be different in
economic growth also continue to rise, we will one such places, in which the conifers are likely to act as
day have to use it all, and there will be no room for keystone species. Research is urgently needed where
vegetation not in the service of humanity. If every- logging and deforestation threaten the existence of
one in the world today enjoyed a lifestyle similar to such tropical conifers with local abundancy, e.g. the
the one people have in Europe and the United States species of Agathis in Borneo and Afrocarpus in east-
we would need at least three planets to sustain our- ern Africa. Conifers are also important in nature
selves; with a world population double that of today, because all trees are providers of ecological services
we would need six planets. Edward Wilson, a lead- to a more stable environment. Climate regulation is
ing writer on evolution and extinction, has put the one; watershed protection preventing erosion and
ecological absurdity of our numbers succinctly, not- assuring a steady supply of clean water in streams
ing that we are now 100 times more numerous than are other important functions of forests. In valleys
the most abundant species of large animals that has with human habitation, the prevention or limitation
ever existed. There is no way that we can draw upon of the effects of snow avalanches in mountainous
the resources of the planet to such a degree without areas is obviously important. On slopes where ava-
drastically reducing the state of most other species. lanche stopping conifer forests have been removed
(Wilson, 1992). That state for most species means it is nearly impossible to restore them and in popu-
extinction. The prospect that within a few decades lated valleys artificial avalanche barriers will have to
far more people will live in cities than in the coun- be erected. They are expensive and not as effective
tryside (the 5050 balance line has recently been as the natural conifer forests. That conifers provide
crossed) threatens to detach humanity further from these services in many parts of the world is the logi-
its biological resources, with dire consequences. cal corollary of their dominant distribution in the
northern hemisphere. The environmental services known. And finally, we would have to recommend
of natural conifer forests, coupled with the creation action and work to ensure these conservation mea-
of habitat to numerous species of animals, fungi and sures are implemented and maintained. The assess-
plants and the provision of material and immaterial ment of conservation status of species on a global
goods to human society are increasingly understood scale is largely undertaken by the International Union
as vital. The proper management of these forests for the Conservation of Nature (IUCN). This Non-
aims at the perpetuation of the original ecosystem Governmental Organisation (NGO), based in Gland,
on large spatial and temporal scales and at least in Switzerland but operating worldwide, works on this
the temperate climate zones of the world utilization through its Species Survival Commission (SSC)
is moving in the direction of such true sustainability. and numerous Specialist Groups, made up of biolo-
32 However, recent Food and Agriculture Organisation gists and other specialists with special knowledge of
of the United Nations (FAO) figures are showing certain groups of animals, plants, or other defined
that subtropical and tropical coniferous forests are groups. The Conifer Specialist Group is responsible
declining faster than any other forest types. Outside for conifers in this context; it has a membership of
a limited number of reserves, exploitation not utili- around 40 individuals and is currently chaired by the
zation is still the rule in much of the tropics. Intrinsic author of this Handbook. The conservation status of
values to conifers may provide incentives to make a species involves knowledge of the extent of occur-
sure they do not become extinct due to our activi- rence (EOO) in nature (excluding all introduced,
ties. These can be aesthetic, cultural, emotional and planted and naturalized occurrences) and the area
scientific. People in all cultures show a remarkable of occupancy (AOO) within the natural range, both
veneration for individual trees, especially if these calculated in square kilometers. These two factors
are old, large, or grow in special places. Thomas determine how widespread (EOO) and how com-
Pakenham, writer, photographer and planter of mon (AOO) a species is. Population size is measured
trees on his estate in Ireland, gives many examples as numbers of mature (reproducing) individuals in
from many countries in his books on remarkable total, or in subpopulations when occurring in scat-
trees, quite a number of which are conifers. The tered or fragmented groups of individuals. Counting
popularity of his books, illustrated with his similarly individuals of trees is only practicable if there are not
remarkable photographs, amply demonstrates the too many, so many estimates are based on EOO and
fascination people still have with large and ancient AOO plus numbers of subpopulations. The conserva-
individual trees. Many conifer species are apparent tion status is also influenced by trends in the popula-
relicts of a geological past when they, or their clos- tion: are the numbers of individuals declining, stable
est relatives, were still abundant. When introduced or increasing? Past, present, as well as projected future
by man to other continents, sometimes returning to trends are of concern and can be used in a conser-
where they once grew, some have done well. Perhaps vation assessment. IUCN has developed criteria for
therein lies the main reason to conserve them, it conservation assessment of this kind, which gives
offers the opportunity to restore something of a lost a species a rating on a categorical scale of conser-
world. There is also an obvious scientific interest, a vation status. An assessed and evaluated species is
curiosity value if you like, in these distinct species then added to the IUCN Red List with its category
representing the past. Given the threat of extinction of threat and the criteria used. The development of
to so many species, prioritization of the species in these criteria has taken considerable time because a
need of conservation may be necessary to save at single system had to be made to work for all organ-
least some of them. Phylogenetic distinction might isms. As a result, there have been two versions (dated
be a criterion that is well worth using to shortlist 1994 and 2001) which are currently both valid in
species for conservation action. terms of having added species to the Red List. The
Conifer Specialist Group assessed most of the (now)
The first task before us if we want to address the 614 recognized conifer species in the period between
increasing threats of extinction of conifer species is to 1994 and 2001 and only about 100 since then, some
assess the conservation status of these species. Types of the latter a second time. These assessments are
of threat need to be identified, with their causes if cited in this book. Re-assessment of the ca. 500 earlier
assessments is urgent, but was still in progress at the change as species are re-assessed, partly due to more
time of publication of this Handbook. The categories and better data, but also due to real changes in their
are now as follows: status. If these two causes are separated, it will be
possible to see a trend in the overall conservation
Extinct (EX) status of conifer diversity. This will then result in a
Extinct in the Wild (EW) Red List Index for conifers, which will be a measure
Critically Endangered (CR) of success, or failure, of conservation action because
Endangered (EN) it will tell us in what direction this diversity is mov-
Vulnerable (VU) ing. It is very important that several non-related
Near Threatened (NT) taxonomic groups of organisms with worldwide dis-
Least Concern (LC) tributions, geographically aa well as ecologically, are 33
Data Deficient (DD) being assessed as completely as possible and as soon
Not Evaluated (NE) as possible. The Convention on Biological Diversity
(CBD) has recommended to develop a number of
The abbreviations are used with all species, subspe- indicators, including one based on changes in status
cies, and varieties treated in this Handbook. The of threatened species, to be able to monitor prog-
coded (abbreviated) criteria are also given, but their ress towards the target of reducing the rate of loss
full explanation would be too extensive to repeat of biological diversity (Baillie et al., 2004). IUCN
here and the reader is referred to the originals has therefore developed Red List Indices for two
(IUCN, 1994, 2001). Taxa assessed and evaluated to such groups: Birds and Amphibians. More taxo-
be in any of the three categories printed in bold are nomic groups with repeated complete assessments
the taxa threatened with extinction in a foreseeable are urgently needed in order to provide data for the
future if trends of decline continue and if no ade- investigation of trends in the conservation status of
quate measures are taken to alleviate the situation. biodiversity. The only taxonomic groups of plants
As calculated when writing this in April 2009, 230 of with suitable baseline data at present (2009) are the
794 accepted taxa (species, subspecies and varieties) Cycads and the Conifers. Conifers are more valuable
or 29% were threatened with extinction. It is slightly for this purpose than Cycads, of which the distribu-
more complicated to calculate how many species tion, although wide, is much more restricted both in
would fall in this broad category, because a species terms of geography and of ecology. Conifers occur
may have one variety at risk and another variety not indeed worldwide and occupy nearly all the major
at risk. The categories of threat are therefore assigned biomes of the terrestrial world. It is hoped that the
to the lowest rank in case of a subdivided species. attention this Handbook may bring to the plight of
Only if all varieties recognized in a species are at risk, conifers in the world will help to put adequate and
the entire species is at risk. This is true for ca. 180 or sufficient conservation policies and action in place
30% of all conifer species. These figures are likely to to indeed slow down their extinction rate.
SY N O P SI S O F FA M I L I E S A N D G E N E R A
This synopsis gives brief character sketches of taxa Agathis Salisb.
in the ranks of genus to family. Only those char-
acters are mentioned which together would suffice Monoecious trees. Leaves subopposite to opposite,
to identify the taxon; these are not necessarily the leaf lamina broad and flat, with distinct petiole. Pollen
same for all taxa, and comparisons between them cones solitary, relatively small. Seed cone scales with 35
are therefore limited to those necessary for identi- an imbricate, rounded margin. Seeds with a single
fication. In this Handbook 8 families and 70 gen- broad wing, becoming detached from the scale.
era are recognized within the order Coniferales, or
conifers. The families are here given in alphabetical Wollemia W. G. Jones et al.
order, as no satisfactory classification at this level
based on all necessary evidence seems possible Monoecious trees. Leaves helically attached, sessile,
(Farjon, 2007, 2008). The hypothetical relation- adult leaves arranged in 4 rows (tetrastichous), linear.
ships of genera within families are indicated by Pollen and seed cones terminal on primary branches,
their groupings; some of these groupings have been relatively small. Seed cone scales with an apical free
named at the rank of subfamily, others are here extension. Seeds circumferentially winged, remain-
presented as informal and separated from each ing attached to the scale; wing(s) narrow.
other by a double space only. The relationships in
Podocarpaceae are most tentative, as they are based Araucaria Juss.
on phylogenies derived from recent cladistic analy-
ses of molecular and morphological data, which Dioecious or monoecious trees. Leaves in helical
are not in agreement for several clades. The DNA- arrangement, sessile, lamina broad and flat, or scale-
based analyses tend to place Phyllocladaceae within like, persistent on falling branches. Pollen cones soli-
Podocarpaceae. Perhaps it could be assumed that tary or in small clusters, relatively small to very large.
the former arose from the latter. The names of fam- Seed cone scales with an apical free extension. Seeds
ilies and genera are given with their abbreviated without wings, remaining attached to the scale.
authorities following Brummitt & Powell (1992).
For more detailed recent classifications down to
species see for Cupressaceae Farjon (2005a), for Cephalotaxaceae Neger
Pinaceae excluding Pinus Farjon (1990), for Pinus
Richardson (ed., 1998) and Farjon (2005b) and for Cephalotaxus Siebold & Zucc. ex Endl.
Taxaceae Cope (1998).
Dioecious evergreen shrubs or small trees. Leaves
pectinately arranged becoming subopposite, linear-
Araucariaceae Henkel & W. Hochst. lanceolate, with two prominent stomatal bands on
the abaxial side. Pollen cones aggregated in capitu-
Dioecious or monoecious evergreen, highly resin- lae. Seed cones reduced, with opposite fertile bracts.
ous trees. Tree architecture according to Massarts Seeds 12 per cone, exposed, large, completely sur-
and Rauhs models. Leaves in helical arrangement rounded by a fleshy aril.
or subopposite to opposite; lamina broad and flat,
or scale-like. Pollen cones catkin-like, sometimes
large. Seed cones large, globose, mostly disinte- Cupressaceae Gray
grating. Seed cone scales predominantly consisting
of the bract, but with a fused seed scale bearing a Aromatic, evergreen or deciduous, monoecious or
single seed. dioecious shrubs or trees. Leaves on (pen)ultimate
branchlets linear, needle- or scale-like, spirally Sequoioideae Saxton
arranged, ternate or decussate (rarely quadrate) in
mature plants. Pollen cones small, terminal, rarely Deciduous or evergreen, monoecious trees. Leaves
axillary, solitary or sometimes clustered in groups helically arranged or opposite, linear or scale-like.
of 27. Seed cones terminal, simple or semi-com- Pollen cones solitary or numerous in spike-like
pound, globose, ovoid or conical. Seed cone scales shoot systems. Seed cones barrel-shaped or ovoid,
consisting of transformed bracts, true seed scales with peltate bract-scale complexes. Seeds axillary to
absent or sometimes rudimentary. Seeds 1-many cone scales, with 2 marginal wings.
axillary to each cone scale, with or without wings.
Metasequoia Hu &W. C. Cheng
36
Cunninghamioideae (Zucc. ex Endl.) Quinn Deciduous, monoecious trees, dropping foliage
branchlets, not individual leaves. Leaves opposite,
Cunninghamia R. Br. linear, spreading at nearly right angles to the shoot.
Pollen cones numerous in spike-like shoot systems.
Evergreen, monoecious trees, capacity to coppice Seed cones terminal on 25cm long, scale-leaved
profound. Leaves helically arranged, linear-lanceo- shoots, subglobose, barrel-shaped or fusiform. Seeds
late; leaf margin serrulate. Pollen cones numerous in numerous, with 2 marginal wings.
clusters. Seed cones subterminal, persistent (falling
with foliage branches), with thin, coriaceous scales. Sequoia Endl.
Seeds 23 per fertile scale, with 2 marginal wings
1mm wide. Evergreen, monoecious (very tall) trees, often
sprouting from lignotubers. Leaves alternate, mostly
linear, pectinate on shaded shoots. Pollen cones on
Taiwanioideae L. C. Li the same branches as seed cones, solitary. Seed cones
terminal on short branchlets, more or less ovoid,
Taiwania Hayata 1530mm long. Bract-scale complexes helically
arranged, parting to release the marginally winged
Evergreen, monoecious trees. Leaves helically seeds.
arranged, imbricate, in young trees falcate-subulate,
ultimately scale-like. Pollen cones in terminal clus- Sequoiadendron J. Buchholz
ters on branchlets with scale leaves. Seed cones ter-
minal, solitary, small, with thin, coriaceous scales. Evergreen, monoecious (giant) trees. Leaves heli-
Seeds usually 2 per fertile scale, each with 2 small cally arranged in 3 ranks, imbricate, scale-like.
wings. Pollen cones on the same branches as seed cones but
well above them, solitary. Seed cones terminal on
short branchlets, 3070(95) cm long. Bract-scale
Athrotaxoideae L. C. Li complexes helically arranged, parting to release the
flattened, unequally winged seeds.
Athrotaxis D. Don
Evergreen, monoecious trees. Leaves decussate, Evergreen, monoecious trees. Leaves scale-like
imbricate, scale-like, (strongly) dimorphic, decur- (adult) and acicular-linear (juvenile), in one species
rent. Pollen cones solitary, subglobose to oblong. both occur in mature trees; adult leaves decussate,
Seed cones subglobose; bract-scale complexes dimorphic, juvenile leaves in whorls of 4, monomor-
decussate, spreading at maturity. Seeds with 2 very phic. Pollen cones solitary, yellow. Seed cones open-
unequal wings. ing wide; bract-scale complexes 4 (or 6) in decussate
pairs. Seeds flattened, with 2 lateral wings.
Juniperus L.
38 Austrocedrus Florin &Boutelje
Evergreen, monoecious or dioecious shrubs or trees.
Leaves decussate or ternate, scale-like or acicular, Evergreen, dioecious shrubs or trees. Leaves decus-
decurrent or articulate. Pollen cones terminal or sate, in opposite ranks, scale-like, dimorphic; facial
axillary. Seed cones terminal on axillary (dwarf) leaves very small. Pollen cones solitary, small.
shoots, globose, semiglobose or ovoid, indehiscent, Seed cones on ultimate branchlets with non-mod-
usually red, glaucous or blue; bract-scale complexes ified leaves, ovoid-oblong; bract-scale complexes
in 14 decussate or ternate whorls, entirely fused, 4, decussate, the upper scales twice as large as the
usually soft. Seeds 18 per cone, wingless. lower. Seeds with 2 wings, one rudimentary, the
other well developed.
Microbiota Kom.
Libocedrus Endl.
Evergreen, monoecious, decumbent shrubs. Leaves
decussate, imbricate, scale-like, weakly dimorphic, Evergreen, monoecious shrubs or trees. Leaves
turning copper-brown or purplish brown in winter. decussate, imbricate, scale-like, decurrent, strongly
Pollen cones axillary. Seed cones hidden in shoot dimorphic. Seed cones terminal on flattened or
axils, minute, reduced to 2 pairs of bract-scales, the quadrangular branchlets, subtended by 45 decus-
upper enclosing a single, ovoid, wingless seed. sate, transitional leaf pairs; bract-scale complexes
in 2 decussate pairs, the upper fertile pair of scales
spreading at maturity, with a long subapical bract
Platycladus Spach tip. Seeds 14 per cone, with 2 very unequal wings.
Evergreen, dioecious trees. Leaves in alternate Monoecious evergreen or deciduous, resinous trees
near-whorls of 3, imbricate, scale-like, decurrent. or shrubs. Leaves spirally inserted on long or short
Seed cones formed by 23 whorls of slightly modi- shoots and solitary or arranged in fascicles of 18
fied scale leaves, followed by 2 alternate whorls of surrounded by a sheath on dwarf shoots (Pinus),
3 fertile, wide spreading scales; columella variably acicular-linear to long linear. Pollen cones often
shaped, trigonal to tripartite. Seeds with 23 narrow grouped close together on long shoots, axillary,
wings. solitary or clustered from a single bud, catkin-like,
deciduous. Seed cones compound, small to very Laricoideae Melchior &Werdermann
large. Bracts free, well developed or rudimentary.
Seed scales persistent or deciduous, woody; bearing Evergreen or deciduous trees. Leaves on long shoots
2 inverted seeds. Seeds with 1 wing or wingless. and/or short shoots, linear. Pollen cones lateral in
leaf axils, or terminal and clustered on short shoots.
Seed cones pendant or erect, deciduous or falling
Pinoideae Pilg. attached to branches. Bracts well developed, often
exserted. Seed scales thin woody. Seeds winged;
Evergreen trees or shrubs. Leaves spirally inserted wing 2 length of seed, adnate.
on long shoots or in fascicles of 18 on dwarf shoots
40 (Pinus), acicular or (long) linear. Pollen cones Larix Mill.
grouped close together on long shoots (Pinus) or
solitary in leaf axils. Bracts of seed cones small but Deciduous trees with pronounced shoot dimor-
conspicuous or rudimentary. Seed scales persis- phism. Leaves on long shoots but predominantly on
tent, thin or thick woody. Seeds winged or wingless. short shoots, linear, lax. Pollen cones terminal and
Number of cotyledons in seedlings high (420). clustered on short shoots. Seed cones remaining
erect, persistent and separated from the tree while
Pinus L. still attached to short shoots and branches. Bracts
exserted or included, not or only slightly longer than
Trees or shrubs. Leaves in fascicles of 18 (com- seed scales.
monly 2, 3 or 5) on dwarf shoots, long linear. Pollen
cones grouped close together in helical arrangement Pseudotsuga Carrire
on long shoots. Seed cones at base of new shoots,
small to very large, persistent or deciduous. Bracts Evergreen trees. Leaves on long shoots, leaving small
rudimentary. Seed scales usually thick woody, some- scars when falling. Pollen cones solitary in axils of
times remaining soft, with a distinct apophysis and leaves. Seed cones becoming pendant at maturity,
umbo. Seeds winged or wingless, wing adnate or falling entire from branches with the peduncle
easily detached. attached. Bracts exserted, longer than seed scales,
with trilobate apex. Seed scales with a rounded
Cathaya Chun &Kuang upper margin. Seeds winged; wing short.
* The genus Cedrus has come out as basal (sister Podocarpaceae Endl.
group) to all other genera of the Pinaceae in some
recent DNA-based cladistic analyses; morphological Dioecious or sometimes monoecious, evergreen
evidence however places it firmly within Abietoideae. trees or (dwarf) shrubs (one species parasitic on
A basal position is not corroborated by the fossil another member of the family). Leaves helically
record. Its position is tentative. arranged, rarely decussate (Microcachrys), appressed
and imbricate to spreading and remote, shapes highly
Keteleeria Carrire variable, ranging from small, appressed scale leaves,
via thin acicular leaves, to dorsiventrally flattened,
Evergreen trees. Leaves on long shoots, broadly lin- linear-lanceolate to broadly lanceolate, large leaves
ear to lanceolate-linear. Pollen cones solitary in leaf (up to 34 9.5cm). Pollen cones mostly simple,
axils. Seed cones remaining erect, breaking up by axillary or terminal, solitary or clustered. Seed cones
disintegration of the rachis. Bracts well developed, axillary or terminal, solitary, mostly much reduced
often exserted. Seed scales persistent. Seeds with and often swelling to form a succulent receptacle;
broad wing dispersed separately; seed germination fertile bracts 1-many. Seeds of most species sur-
hypogeal. rounded by a coriaceous or succulent epimatium.
Evergreen trees with weakly developed shoot dimor- Monoecious trees. Leaves pectinate, short decurrent,
phism. Leaves linear. Pollen cones in umbellate clus- linear or falcate. Pollen cones axillary, solitary or
ters from a single bud. Seed cones remaining more sometimes in pairs, 46mm long. Seed cones termi-
or less erect, falling entire or disintegrating. Seed nal, globular, 912mm diam. Cone scales 1520 per
scales persistent. Seeds with small wing dispersed cone, spirally arranged, swelling at maturity. Seeds 1
separately. per scale (from 2 inverted ovules), enclosed.
Deciduous trees with pronounced shoot dimorphism. Dioecious or rarely monoecious shrubs or more
Leaves on long shoots but predominantly on short commonly trees. Leaves relatively broad, usually
shoots, linear, lax. Pollen cones in umbellate clusters linear-lanceolate or linear-elliptic, coriaceous, with
from the apex of a short shoot. Seed cones termi- a single raised, flat or sunken midrib and stomata
nal on short shoots, remaining erect, disintegrating. in two broad bands on the abaxial side. Pollen
cones axillary, solitary or clustered, catkin-like. Dacrycarpus (J. J. Bennett) de Laub.
Seed cones swelling to form a smooth, succulent,
coloured receptacle. Seeds 1(2) per cone, com- Dioecious or rarely monoecious shrubs or trees.
pletely exposed, drupe-like, completely covered by Leaves trimorphic, with small scale leaves, acicu-
a fleshy epimatium. lar leaves and flattened, linear-falcate leaves. Pollen
cones single or in pairs on axillary short shoots,
Retrophyllum C. N. Page cylindrical. Seed cones solitary, when mature form-
ing an irregular, fleshy and verrucose receptacle.
Dioecious, dwarfed to large trees. Leaves spirally Seeds single, enclosed by a fleshy epimatium.
inserted, lanceolate to narrowly ovate, obliquely
42 directed into subopposite and decussate appar- Dacrydium Sol. ex G. Forst.
ent pairs by twisted petioles in opposite directions,
amphistomatic. Pollen cones axillary or terminal. Dioecious or rarely monoecious shrubs or trees.
Seed cones rudimentary, with a single large seed Leaves dimorphic, scale-like and subulate or acicu-
covered by a fleshy, drupe-like epimatium. lar. Pollen cones small, cylindrical. Seed cones small,
forming a red receptacle. Seeds 12(3) per cone,
Afrocarpus (J. Buchholz & N. E. Gray) becoming more or less erect, protruding from a cup-
C. N. Page like epimatium covering basal part of seed only, lus-
trous brown or nearly black.
Dioecious trees. Leaves twisted in opposite direc-
tions, narrowly lanceolate-elliptic to linear-lanceo- Falcatifolium de Laub.
late, coriaceous, with a single midrib, amphistomatic.
Pollen cones axillary, solitary or in groups of 23, Dioecious shrubs or trees. Leaves dimorphic, with
catkin-like. Seed cones much reduced, not trans- scale leaves on leading and fertile shoots, alternating
formed into a receptacle. Seeds 1 per cone, sub- with more or less distichously spreading, bilaterally
tended by small, withering scales, entirely enclosed flattened, obliquely lanceolate-falcate leaves on veg-
by a swollen, drupe-like epimatium. etative shoots. Pollen cones cylindrical and catkin-
like. Seed cones forming an irregular, red receptacle.
Nageia Gaertn. Seeds erect, surrounded at the base by a swollen epi-
matium, with two lateral ridges.
Dioecious or monoecious trees, rarely shrubs.
Leaves large, flat, broadly ovate-elliptic to lanceolate, Halocarpus Quinn
lacking a midrib and with many parallel, converging
veins. Pollen cones single or in small, spicate groups Dioecious shrubs or trees. Leaves dimorphic, radi-
of 26 on axillary peduncles, ovoid-cylindric. Seed ally spreading, lanceolate-linear to linear ones alter-
cones much reduced, or forming a weakly developed, nating with appressed, rhombic scale leaves. Pollen
fleshy receptacle. Seeds 1 per cone, exposed, entirely cones solitary or with 23 together at the apex of
enclosed by a swollen, drupe-like epimatium. scale-leaved branchlets, small. Seed cones reduced
to a few reddish bracts at the apex of scale-leaved
Acmopyle Pilg. branchlets. Seeds 15 per cone, erect, at base sur-
rounded by a swollen epimatium forming a white or
Dioecious (or sometimes monoecious?) small trees. yellow collar, lustrous black.
Leaves dimorphic, small and scale-like on leading
and fertile shoots, larger and falcate-linear on lateral, Lagarostrobos Quinn
vegetative shoots. Pollen cones cylindrical, more
or less erect. Seed cones solitary or occasionally Predominantly dioecious trees, with lower branches
grouped, when mature forming an irregular, fleshy frequently layering. Adult leaves imbricate and
and verrucose receptacle. Seeds single, enclosed by appressed, rhomboid in appearance, 11.5 1mm.
a fleshy epimatium. Pollen cones terminal, sessile. Seed cones terminal
on decurved short branchlets, 45mm long. Seeds Pherosphaera W. Archer
up to 58 per cone, usually fewer, light brown,
enclosed at base by a dry, papery epimatium. Dioecious shrubs. Adult and juvenile leaves simi-
lar, spirally arranged, 24(6) mm long and mostly
Lepidothamnus Phil. scale-like. Pollen cones terminal, globose to ovoid.
Seed cones terminal, with (3)58 spreading, fertile
Dioecious or sometimes monoecious, creeping or scales, not fleshy at maturity. Seeds usually 14 per
erect shrubs or trees. Leaves dimorphic, with lin- cone, exposed, erect.
ear, spreading juvenile ones usually gradually giving
way to appressed, ovate-rhombic and gibbous scale Prumnopitys Phil.
leaves. Pollen cones terminal or sometimes lateral. 43
Seed cones terminal, solitary, very small, consisting Dioecious trees or shrubs. Leaves small, in lateral
of 35 yellow or reddish bracts. Seeds 1 per cone, shoots pectinately arranged, flattened, falcate-linear.
purplish brown or black, surrounded at base by a Pollen cones aggregate on relatively long, leafless
membranous epimatium. branchlets, cylindrical. Seed cones reduced to a few
bracts. Seeds enclosed in a drupe-like, fleshy, yellow,
Manoao Molloy red or black epimatium.
Predominantly dioecious trees; sucker shoots from Sundacarpus (J. Buchholz & N. E. Gray)
horizontal underground stems prolific. Adult leaves C. N. Page
imbricate and appressed, rhomboid in appearance,
11.5 1mm. Pollen cones terminal, sessile. Seed Dioecious trees. Adult leaves broadly linear, 515cm
cones terminal on distally curved short branchlets, long, 615mm wide, petiolate, with a single mid-
34mm long. Seeds 13(-5) per cone, dark purple vein. Pollen cones axillary, often 37 on a stalk,
to black, the basal half or more enclosed by a swol- cylindrical. Seed cones much reduced, not forming
len, fleshy epimatium. a receptacle. Seeds 1 per cone, large, covered by a
glaucous purple, fleshy but firm epimatium.
Microcachrys Hook. f.
Sciadopityaceae Luerss.
Monoecious or (temporarily) dioecious, prostrate
shrubs. Leaves on lateral branchlets decussate, Sciadopitys Siebold &Zucc.
imbricate, appressed, short triangular in appearance,
11.5 1mm. Pollen cones terminal, more or less Evergreen, monoecious trees. Foliage consisting
recurved. Seed cones terminal, with ca. 20 helically of linear cladodes (needles) in pseudo-whorls on
arranged, fleshy, red bracts. Seeds 1 per fertile bract shortened terminal sections of long shoots. True
scale, at base partly covered by an epimatium. leaves much reduced, scale-like. Pollen cones in ter-
minal or sublateral clusters. Seed cones terminal or
sublateral, ovoid to cylindrical, often exuding resin.
Parasitaxus de Laub. Bract-scale complexes helically arranged, consist-
ing of a partly fused bract and exceeding seed scale.
Small erect monoecious shrubs, parasitic on the Seeds basal to seed scales, inverted, with 2 wings.
roots of Falcatifolium taxoides. Leaves spirally
arranged, imbricate, scale-like, decurrent, reddish.
Pollen cones usually on the same branching systems Taxaceae Gray
as seed cones, terminal, solitary. Seed cones reduced
to a few bracts with a single seed surrounded by a Dioecious or rarely monoecious evergreen shrubs or
hard, glaucous white epimatium. trees. Leaves helically or decussately inserted, usually
pectinately arranged, linear to lanceolate with a almost entirely enclosing the seed and only leaving
single midrib, hypostomatic, with two bands of sto- the seed apex free, ovoid, smooth, at first glaucous
mata. Pollen cones axillary to foliage leaves and soli- green ripening to orange.
tary or aggregated in umbellate clusters of racemes.
Seed bearing structures consisting of axillary dwarf Pseudotaxus W. C. Cheng
shoots with terminal, erect ovules. Seeds surrounded
by a fleshy or succulent aril; aril partially or entirely Dioecious shrubs. Leaves helically arranged, dis-
enclosing the seed, becoming red, purple or yellow tichous, spreading, linear. Pollen cones axillary,
when ripe. solitary. Seed aril cupular, only partly enclosing the
seed, succulent and white when ripe.
44 Amentotaxus Pilg.
Taxus L.
Dioecious shrubs or small trees. Leaves usually oppo-
site-decussate, pectinately arranged by twisted peti- Dioecious shrubs or trees. Leaves helically inserted,
olate leaf bases, linear-lanceolate, straight or slightly twisted at petiolate base and becoming pectinate or
falcate, large. Pollen cones aggregated in umbellate distichous, linear or falcate. Pollen cones axillary,
clusters of racemes. Seed-bearing structures usually solitary. Seed aril cupular, only partly enclosing the
in groups near the apex of vegetative shoots. Aril seed, succulent and red, orange or yellow when ripe.
surrounding the ripe seed greatly enlarged, ellipsoid
or ovoid, fleshy and succulent, red or purple, com- Torreya Arn.
pletely hiding the small seed.
Dioecious or rarely monoecious trees or some-
Austrotaxus R. H. Compton times shrubs. Leaves helically inserted, pectinately
arranged, linear-lanceolate, rigidly coriaceous, acu-
Dioecious trees. Leaves alternate (helical), spreading minate or pungent. Pollen cones axillary, solitary.
forward at 3060 to the shoot axis, narrowly lan- Aril almost completely surrounding the large seed,
ceolate, large. Pollen cones axillary, solitary. Seed aril fleshy, purplish green to bluish black when ripe.
TA XO N OM IC T R E AT M E N T O F FA M I L I E S
Key to the families development); seeds 1-many, axillary or on the
base of each bract (rarely only a single seed per
This Handbook recognizes eight families within the cone) 4
conifers (currently with 70 genera and 614 species); 3b. Seed cones either with much reduced, obscure
some of their diagnostic characters are here used scales (whether bracts or seed scales, in a few 45
in the key to families. If the family to which a spe- instances with many very small bracts) or lack-
cies has been assigned is not known to the user, it ing any kind of scales and reduced to a single,
is necessary to start with this key and then proceed terminal seed with a surrounding epimatium
to the key to genera under the family determined. or arillus 5
If the family is known, one can proceed directly to 4a. True leaves scale-like or acicular (needle-like),
the latter key to determine the genus. Keys to spe- cataphylls absent, phyllodes absent
cies within genera are provided under each genus; Cupressaceae
the species are arranged in alphabetical order (A-Z) 4b. True leaves reduced to tiny cataphylls (usually
throughout using the Latin binomials, which enables brown scales); phyllodes (pseudo-leaves) in the
the user to find the genus determined using the keys axils of these green, needle-like, growing rhyth-
with ease. mically in pseudo-whorls on shoots
Sciadopityaceae
1a. Seed cones with seed scales in the axils of bracts 5a. Seed cones reduced to a single seed; seed ter-
(bracts can be much reduced in mature, woody minally placed at a scaly dwarf shoot, partly or
cones but are conspicuous in immature cones at completely surrounded by a succulent or fleshy
early stages of development); i.e. cones clearly aril Taxaceae
compound and never reduced; seeds two on the 5b. Seed cones apparent, with much reduced,
adaxial (upper) side of each fertile scale; adult obscure scales (usually bracts; if these are seem-
green leaves acicular-linear Pinaceae ingly absent, seed on a stout or fleshy pedun-
1b. Seed cones with seed scales fused with bracts cle) or in a few instances with many, very small
(bracts make up the bulk of the cone), or with bracts 6
bracts only (which may be much enlarged, 6a. Pollen cones aggregated in more or less globose
swollen and/or woody at maturity), or with capitulae (heads); bracts of (reduced) seed
scales obscure, much reduced or absent; seeds cones decussate Cephalotaxaceae
either single or more than two per fertile scale; 6b. Pollen cones solitary or clustered but not in glo-
adult green leaves scale-like, or acicular, or with bose capitulae; bracts of (reduced) seed cones
a distinct lamina, or replaced by phylloclades alternate or helically arranged (but sometimes
(phyllodes) 2 seemingly absent being enveloped by a swollen
2a. Seed cones with seed scales fused with bracts, receptacle) 7
or with bracts only which are much enlarged 7a. True leaves scale-like, acicular or with a distinct
and often swollen at maturity and may then lamina, always simple; cataphylls absent; phyl-
form a compact, globose cone 3 loclades absent; cones arising in the axils of
2b. Seed cones with seed scales fused with bracts; leaves Podocarpaceae
bracts forming the largest part of the cone 17b. True leaves reduced to tiny cataphylls, usually
scales; a single inverted seed per scale soon deciduous; phylloclades (pseudo-leaves)
Araucariaceae leaf-like, green, simple or compound; cones
3a. Seed cones consisting of bracts forming the arising on the edges of phylloclades or on sepa-
cone scales only (sometimes with rudimentary rate determinate shoots (axillary to bracts)
seed scales only visible at very early stages of Phyllocladaceae
Descriptions of families and keys to genera Three genera: Agathis (17 spp.), Araucaria (19 spp.)
and Wollemia (1 sp.); total 37 species.
Araucariaceae Henkel & W. Hochst., Syn.
Nadelhlz.: xvii, 1. 1865. (nom. cons.). Type: Distribution
Araucaria Juss.
Malesia: all major islands except Jawa and Lesser
Description Sunda Islands; Australia: New South Wales,
Queensland. SW Pacific: New Caledonia, Vanuatu,
Dioecious or monoecious evergreen, highly resin- Fiji, Norfolk Island, New Zealand (North Island).
ous trees. Tree architecture according to Massarts South America: SE Brazil, NE Argentina; S Chile,
46 and Rauhs models. Resin canals in bark, leaves and SW Argentina (Andes).
seed cones. Bark hard and smooth, exfoliating with
rounded or irregular flakes (Agathis), rough and Key to the genera
exfoliating in horizontal strips and eventually deeply
fissured (Araucaria), or forming many irregular 1a. Bark hard and smooth, exfoliating with
pustules (Wollemia). Branches in rhytmic pseudow- rounded or irregular flakes; leaves distinctly
horls, spreading and plagiotropic (Massarts model) petiolate; seed cones globose, with imbricate
or ascending to become orthotropic (Rauhs model), scales lacking an extended apex Agathis
sometimes profusely reiterating or sprouting from 1b. Bark rough, exfoliating in horizontal strips or
dormant buds in roots near the base of stems. Foliage forming many irregular pustules; leaves ses-
branchlets with or without terminal buds. Leaves spi- sile; seed cones ovoid-globose to globose; cone
rally arranged or subopposite, scale-like and adnate scales with an extended, free apex 2
or laminar and sessile or short petiolate, imbri- 2a. Bark forming many irregular pustules in
cately covering the shoot or free and more or less mature trees; leaves opposite or subopposite,
distichously spreading, sometimes forming 4 ranks mostly linear, distichous or tetrastichous (in 2
(Wollemia), more or less coriaceous, with numerous or 4 ranks); pollen and seed cones terminal on
parallel veins originating from basal dichotomies first-order branches Wollemia
and few to numerous resin canals. Pollen cones axil- 2b. Bark exfoliating in horizontal strips; leaves
lary to leaves, solitary or in small clusters, small or spirally arranged, scale-like or triangular to
large, much elongating after anthesis and becoming lanceolate, mostly equally divided around the
cylindrical; microsporophylls numerous, helically shoot; pollen and seed cones lateral on higher
inserted, crowded, with imbricate or tesselate heads, order foliage branches Araucaria
each with 420 oblong pollen sacs containing non-
saccate pollen. Seed cones terminal on long shoots
(Wollemia) or lateral on short, pedunculate, leafy Cephalotaxaceae Neger, Nadelhlzer: 23, 30. 1907
shoots, solitary, erect, ovoid or subglobose, some- (nom. cons.). Type: Cephalotaxus Siebold & Zucc. ex
times massive (Araucaria bidwillii has the heaviest Endl.
cones of all conifers), usually disintergrating leaving
the rachis on the tree. Bracts helically inserted on the
Description
rachis, much developed, flattened, with a thickened
distal margin and with or without a terminal elon- See the genus description.
gated cusp, forming the bulk of the cone. Seed scales
much reduced, axillary to and almost entirely fused One genus: Cephalotaxus, with 8 species.
with the bract, with or without a small, free apical
ligule, more or less enclosing a single, inverted seed, Distribution
concrescent with the seed scale or free, wingless or
with a single wing or 2 unequal wings. Seedlings with As for the genus.
2, sometimes deeply divided, cotyledons. Number of
chromosomes (diploid) 2n = 26.
Taxonomic Notes until its relationships are unambiguously resolved,
it seems advisable to retain a monogeneric family
The family Cephalotaxaceae has been variously cir- Cephalotaxaceae.
cumscribed by different authors, or included in
the Taxaceae s. l. This situation led Page (1990) to
remark that obscurity is probably the only aspect of Cupressaceae Gray, Nat. Arr. Brit. Pl. 2: 222, 225.
the generic and family affinities.... about which we 1821. (nom. cons.), [Cupressideae]. Type:
can be totally sure. He included Amentotaxus and Cupressus L.
Cephalotaxus in Cephalotaxaceae, but his circum-
scription has not been generally accepted. Florin
Description
(1948) excluded Taxaceae s. str. from true conifers, 47
but included Cephalotaxus, which he considered to Aromatic, resinous, evergreen or sometimes decid-
occupy an isolated position among the recent rep- uous, monoecious or dioecious shrubs or trees,
resentatives of this class. Cephalotaxus appeared ranging from diminutive prostrate shrubs to trees
to Florin to have several ovules aggregated into a exceeding 100 m. Bark fibrous or brittle, exfoliating
cone, while the other genera lacked evidence of a in longitudinal strips or small plates. Branches erect
cone, or the reduction of one. A number of mor- or spreading; foliage branches erect to pendulous,
phological studies of both male and female repro- initially covered with (decurrent) leaf bases until
ductive organs have brought the two groups more secondary growth replaces these with bark, terete,
closely together, often inferring a more primitive (quadr)angular or more or less flattened, orthotropic
state in Cephalotaxus compared with Taxaceae s. or in more or less plagiotropic and often frond-like
str. The male cones in Taxaceae were interpreted sprays. Buds mostly absent; (seasonal) abscission of
by Wilde (1975) as reduced structures derived from (pen)ultimate branchlets, not of single leaves. Foliage
more clearly compound systems in Cephalotaxaceae. leaves simple, usually polymorphic dependent on life
These differences were confirmed in an elaborate phase, with juvenile, transitional and mature forms,
study by Mundry (2000). The structure and ontog- and on growth of foliage branches. Leaves on (pen)
eny of the female reproductive organs however, are ultimate branchlets linear, needle- or scale-like, spi-
essentially similar in Cephalotaxus and Taxaceae. rally arranged, ternate or decussate (rarely quad-
The cone Florin observed in Cephalotaxus results rate) in mature plants, also in whorls of 4 in juvenile
from a swelling of the bracts subtending the ovules, plants and in some mature plants; decussate scale
of which more in a single reproductive unit are leaves pairwise dimorphic on flattened branchlets.
retained, at least to the pollination stage, in this genus Leaf resin duct or cavity mostly single, often with
than in the others. Other similarities are evident, a dorsal gland in scale leaves. Leaf vascular bundle
but there are also some marked differences; Chen & single. Pollen cones small, terminal, rarely axillary,
Wang (1990) found the numbers of free nuclei in the solitary or sometimes clustered in groups of 27, ses-
female gametophyte to be extremely high but vari- sile on foliage branchlets or on dwarf shoots, sim-
able (10244096) in Cephalotaxus, and relatively low ple, deciduous. Microsporophylls spirally arranged,
and constant (256) in 4 of the 5 genera of Taxaceae ternate or decussate in congruence with leaf phyl-
(Torreya was not investigated). Phylogenies recon- lotaxis on the cone-bearing shoot, small and thin,
structed from molecular data (DNA nucleotide (sub)peltate. Pollen sacs (microsporangia) abaxial,
sequences) tend to place Cephalotaxus basal (as free, 210(-14) per microsporophyll, in one or two
sister to) a clade with Taxaceae s. str. (e.g. Cheng rows, longitudinally dehiscent, containing spheroi-
et al., 2000; Quinn et al., 2002), but in the latter dal, non-saccate pollen. Seed cones terminal, soli-
study the consensus tree shows a polytomy within tary or sometimes secondarily clustered, sessile or
a single clade. This can be interpreted as evidence pedunculate, simple or semi-compound, globose,
that Cephalotaxus is best placed within Taxaceae ovoid or conical, deciduous or persistent. Bract-
(Quinn et al., 2002), but the total evidence is still scale complexes spirally arranged, in whorls of 34
ambiguous. A solution of sorts could be to recog- or decussate in congruence with leaf phyllotaxis on
nize this genus as a subfamily Cephalotaxoideae, but the cone-bearing shoot, consisting of transformed
bracts enlarged with subsidiary intercalary growth, phyllotaxis as a distinctive character; at that time
rarely with elements of ovuliferous scales, forming Metasequoia (with opposite phyllotaxis but in other
(semi-)woody or soft, imbricate or valvate, free or characters similar to e.g. Sequoia and Taxodium in
(partially) fused, (sub)peltate, ovate or oblong cone Taxodiaceae) was unknown. Another distinction
scales with or without an abaxially protruding bract claimed in earlier works was the position of ovules in
tip. Ovules at base of bracts, axillary or sometimes developing cones: axillary to bracts (Cupressaceae)
terminal, erect (or secondarily inverted), single to or basal upon them (Taxodiaceae). In reality, as has
numerous. Seeds winged or unwinged, wings 13 now been shown in detailed observations of ontog-
per seed, often unequal in size and shape, derived eny of cones using scanning electron microscopes
from thin or thick and hard seed coat, seed often (SEM), there is no clear cut distinction, but a grade
48 with resin pits. Seedlings with 26(-9) cotyledons; through both families. If there is a line of demar-
germination epigeal. Chromosome number (dip- cation between the two positions, it lies within the
loid) = 22, sometimes tetra- or hexaploid (Sequoia traditional Taxodiaceae (see Farjon, 2005a for an
sempervirens). overview). It was Eckenwalder (1976) who first dem-
onstrated in a comprehensive phenetic analysis of
30 genera: Actinostrobus (3 spp.), Athrotaxis (3 spp.), morphological and other characters, that apart from
Austrocedrus (1 sp.), Callitris (15 spp.), Calocedrus Sciadopitys (formerly classified in Taxodiaceae),
(4 spp.), Chamaecyparis (5 spp.), Cryptomeria (1 sp.), no taxon assigned to Taxodiaceae was separable
Cunninghamia (2 spp.), Cupressus (15 spp.), Diselma from a larger group including taxa classified in the
(1 sp.), Fitzroya (1 sp.), Fokienia (1 sp.), Glyptostrobus Cupressaceae. Phylogenetic analyses, both of mor-
(1 sp.), Juniperus (53 spp.), Libocedrus (5 spp.), phological data and of molecular (DNA) data, and
Metasequoia (1 sp.), Microbiota (1 sp.), Neocallitropsis of combined data, all demonstrate that the genera
(1 sp.), Papuacedrus (1 sp.), Pilgerodendron (1 sp.), of traditional Taxodiaceae constitute not a coherent
Platycladus (1 sp.), Sequoia (1 sp.), Sequoiadendron clade (group), but a grade of taxa with on the whole
(1 sp.), Taiwania (1 sp.), Taxodium (2 spp.), Tetraclinis less advanced character states. These genera are also
(1 sp.), Thuja (5 spp.), Thujopsis (1 sp.), Widdringtonia predominantly monospecific, with in several cases
(4 spp.) and Xanthocyparis (2 spp.); total 135 species. more species known from the fossil record. They
therefore constitute a loose assembly of relict species
Distribution with more or less basal positions in the phylogeny of
the Cupressaceae. It is interesting to note that palaeo-
Cosmopolitan. Northern hemisphere: Macaronesia; botanists, who have been more hesitant than other
N Africa (including one locality in Central Sahara). systematists to abandon the concept of Taxodiaceae,
Eurasia: Europe (including Iceland), SW Asia (excl. are now also beginning to interpret Cupressaceae in
driest deserts), Central Asia, Mongolia, Siberia, the here accepted circumscription (e.g. Stockey et
Russian Far East (including Kamchatka), Hindu al. in Farjon, 2005a; Anderson et al., 2007). This is
Kush, Himalaya, China, Taiwan, Japan, Korea, undoubtedly due to the fact that more fossil remains
Lao PDR, Viet Nam. Australasia: New Guinea, are coming to light that appear to fill in the gaps cre-
Maluku (Moluccas), Australia (including Tasmania). ated by the extinctions.
SW Pacific: New Caledonia, New Zealand. North
America: Canada, S Greenland, USA (including Key to the genera
Alaska), Mexico. Central America S to Honduras.
Caribbean Islands. South America: S Chile and 1a. Leaves linear-lanceolate; margins serrulate;
S. Argentina (Andes) to Tierra del Fuego. seed cones > 15mm wide, composed of heli-
cally arranged, coriaceous scales each with 23
Taxonomic notes seeds Cunninghamia
1b. Leaves of a different shape, entire if linear; seed
The family Cupressaceae here includes the for- cones if with helically arranged coriaceous
mer Taxodiaceae. The initial full description of scales, then <15mm wide and 2 seeds per scale
Taxodiaceae by Warming (1884) included spiral 2
2a. Leaves on lateral, short branchlets all linear, flat 9b. Leaves subulate or scale-like on different
and entire 3 mature trees; seed cones (sub)globose, with
2b. Leaves on lateral, short branchlets at least thickened cone scales; pollen cones solitary
in part scale-like or subulate, or a mixture of Athrotaxis
scale-like and linear leaves present on the same 10a. Seed cones berry-like, soft, indehiscent; cone
mature plant 5 scales (almost) entirely fused; leaves scale-like
3a. Leaves on lateral branchlets and cone scales or acicular-linear Juniperus
decussate, lateral branchlets deciduous 10b. Seed cones not berry-like, (eventually) dehis-
Metasequoia cent; cone scales at least partly integer; leaves
3b. Leaves on lateral branchlets and cone scales on lateral branchlets scale-like, rarely acicular
helically arranged 4 or linear 11 49
4a. Leaves scale-like on leading branches, linear 11a. Leaves on lateral branchlets decussate, dimor-
and flat on lateral branchlets, lateral branchlets phic, with differently shaped facials and laterals
evergreen; all cones near or at the end of foliage 12
branchlets Sequoia 11b. Leaves on lateral branchlets decussate or
4b. Leaves (almost) all linear, flat or short acicu- whorled (34), monomorphic or at least not
lar to subulate in one variety; lateral branch- clearly divided in facials and laterals 23
lets (semi-)deciduous; pollen cones numerous 12a. Seed cones (sub)globose when closed, with
on leafless branchlets; seed cones on thicker cone scales of more or less equal length 13
branches, often aggregate Taxodium 12b. Seed cones oblong when closed, with some
5a. Leaves on lateral foliage branchlets and cone paired cone scales longer than other scales 18
scales all helically arranged 6 13a. Seed cones with 4(-56) subpeltate scales in
5b. Leaves on lateral foliage branchlets and cone decussate pairs 14
scales decussate, opposite or whorled 10 13b. Seed cones with 8 or more peltate or subpeltate
6a. Leaves on lateral foliage branchlets scale-like as scales in decussate pairs 16
well as linear, lateral branchlets deciduous 14a. Leaves weakly dimorphic, appressed; columella
Glyptostrobus in seed cone absent 15
6b. Leaves on lateral foliage branchlets scale-like or 14b. Leaves strongly dimorphic, lateral leaves with
subulate or lanceolate, lateral branchlets ever- free apex, or monomorphic linear leaves pres-
green 7 ent; columella present Xanthocyparis
7a. Leaves on lateral branchlets mostly scale-like; 15a. Small tree or shrub; leaves long decurrent,
seed cones ovoid, > 30 mm long with 30 or adnate to branchlets except for the small apex;
more peltate scales; bark on trunk thick and seed cones woody, > 10mm, with 46 winged
soft fibrous Sequoiadendron seeds Tetraclinis
7b. Leaves on lateral branchlets scale-like, 15b. Decumbent shrub; leaves short decurrent; seed
appressed, or subulate or lanceolate; seed cones cones < 5mm, with a single, wingless seed
< 30mm long with less than 30 scales; bark rel- Microbiota
atively thin and scaly 8 16a. Leaves on lateral branchlets lustrous green
8a. Leaves on lateral branchlets of cone-bearing above, with conspicuous, white stomatal bands
trees all subulate, distinctly keeled; seed cones below; seed cones > 11mm 17
squarrose; cone scales cuneate, with small teeth 16b. Leaves green above, with obscure, green-
above the recurved bract tip Cryptomeria ish white stomatal bands below; seed cones
8b. Leaves on lateral branchlets scale-like, subulate <12mm Chamaecyparis
or lanceolate; seed cone scales without small 17a. Seed cones with 2(-3) pairs of larger scales and
teeth above the bract tip 9 23 pairs of reduced scales; leaves thick, facials
9a. Leaves subulate in younger trees only, eventu- and laterals equally long Thujopsis
ally changing to scale leaves; seed cones ellip- 17b. Seed cones with at least 4 pairs of larger, peltate
soid to cylindrical, with thin cone scales; pollen scales, only the upper 12 pairs reduced; leaves
cones aggregate Taiwania thin, laterals exceeding facials Fokienia
18a. Seed cones with 3 decussate pairs of scales, the 27b. Leaves usually in alternate whorls of 3, scale-
proximal pair reduced, the middle pair spread- like or sometimes acicular (juvenile form); seed
ing and fertile, the long distal pair fused and cones common, with 6, rarely with 8, > 3mm
sterile Calocedrus wide scales 28
18b. Seed cones with 2 or 36 decussate pairs of 28a. Mature type leaves long decurrent, adnate to
scales, the middle and distal pairs spreading branchlets except for the (small) apex; seed
and fertile, or the distal pair reduced and sterile cones with thick woody, valvate scales and a
19 conical, hard columella 29
19a. Seed cones with 2 decussate, fertile pairs of 28b. Mature type leaves short decurrent, loosely
scales 20 appressed or spreading; seed cones with thin
50 19b. Seed cones with 36 decussate pairs of fertile woody, widely spreading scales and a trigonal
and sterile scales 22 to tripartite, soft columella Fitzroya
20a. Seed cone scales with a large bract apex 29a. Valvate, thick cone scales subtended by
Libocedrus appressed, thin sterile scales; leaf apices on ulti-
20b. Seed cone scales with a small bract apex 21 mate branchlets spreading and acute
21a. Facial leaves minute in comparison to laterals Actinostrobus
in all stages; bract apex nearly central on the 29b. Valvate, thick cone scales not subtended by
seed cone scale Papuacedrus sterile scales; leaf apices on ultimate branchlets
21b. Facial leaves slightly smaller than laterals; bract usually appressed or just free. Callitris
apex subapical Austrocedrus
22a. Seed cone scales thin, the distal pair reduced
and connate; seeds winged Thuja Phyllocladaceae Bessey, Nebraska Univ. Stud. 7: 325.
22b. Seed cone scales thickened, the distal pair well 1907. Type: Phyllocladus Rich. ex Mirb.
developed and spreading; seeds unwinged
Platycladus Description
23a. Leaves on lateral branchlets decussate 24
23b. Leaves on lateral branchlets whorled, usually See the genus description.
ternate 27
24a. Seed cones globose to ovoid, with > 6 peltoid One genus: Phyllocladus, with 4 species.
scales; seeds numerous, small, winged
Cupressus Distribution
24b. Seed cones with a different shape and/or fewer
scales; seeds few to numerous, winged or nearly As for the genus.
wingless 25
25a. Shrub to very small tree (to 6 m); seed cones Taxonomic notes
tiny, ca. 3mm long, with 2 pairs of scales; maxi-
mal 2 seeds per cone Diselma The genus Phyllocladus was placed within the
25b. Shrub to tall tree, seed cones > 10 mm long, Podocarpaceae as a subfamily Phyllocladoideae
with 2 or more pairs of scales; > 2 seeds per by Pilger (1903), but elevated to family rank by
cone 26 Bessey (1907), a publication apparently over-
26a. Seed cones subglobose when closed, with thick looked by Keng (1973), who proposed the family
woody scales and numerous seeds per cone; again. Since then, this family status has been con-
leaves on older branchlets more or less spirally troversial. Keng, in a series of papers re-published
arranged Widdringtonia in Taipei (Taiwan) as A monograph of the genus
26b. Seed cones ovoid-oblong when closed, with Phyllocladus (Coniferae) (Keng, 196379), empha-
thin woody scales and 34 seeds per cone; all sised the peculiar phylloclades which function as
leaves decussate Pilgerodendron true leaves and, erroneously, believed they were
27a. Leaves all in alternate whorls of 4, seemingly in homologous with progymnospermous dichotomous
8 rows, lanceolate; seed cones rare, with 8 only branching systems, e.g. those of the Devonian genus
23mm wide scales Neocallitropsis Archaeopteris. We now know that they are adaptive
structures of much more recent origin, unique in formalized way (e.g. cladistic analysis) and more evi-
conifers but evolved independently in some angio- dence is needed to resolve this question to the satis-
sperms. Kengs interpretation was criticised by faction of all.
Quinn (1987) and in a series of subsequent papers
(Chaw et al., 1997; Conran et al., 2000; Quinn et al.,
2002; Quinn & Price, 2003) researchers analysed Pinaceae Spreng. ex F. Rudolphi, Syst. Orb. Veg.: 35.
various DNA sequence data to address the question 1830. Type: Pinus L.
of the phylogenetic relationship of Phyllocladus. The
presented cladograms predominantly show a clade Description
Podocarpaceae consisting of selected representa-
tive taxa, with Phyllocladus in a basal position (sis- Monoecious evergreen or deciduous, resinous trees 51
ter to the other genera). The implication inferred by or shrubs, sometimes to 100 m tall. Resin canals
Quinn and co-workers is that Phyllocladus is a true in wood, bark, leaves and seed cones. Wood with
podocarp despite its highly distinctive morphol- adaxial parenchyma, with normal (non-traumatic)
ogy. If retained in the Podocarpaceae, its phyloge- resin canals and ray tracheids. Branching in rhyt-
netic relationship as inferred from these molecular mic pseudo-whorls on main stem and branches,
data would obviously warrant status as a subfamily. with apical dominance maintained or restored and
Whether one recognizes it instead as a family then limited reiteration (Massarts and Rauhs models);
depends on two criteria. One is the acceptance or branches terminating in seasonally dormant buds.
rejection of paraphyletic taxa: when Phyllocladus Leaves spirally inserted on long or short shoots and
and the Podocarpaceae are recognized in equal solitary, or in fascicles of 18 surrounded by a sheath
rank (both as family) and given the results of these on dwarf shoots (Pinus), acicular-linear to long lin-
analyses (phylogeny represented by cladograms), ear, amphistomatic or hypostomatic, sometimes
the Podocarpaceae would be paraphyletic (i.e. a epistomatic. Leaf anatomy in cross section with 12
group not including all its descendants) instead of vascular bundles encapsulated in a stele and 1-many
monophyletic (including all its descendants). But resin canals variously positioned in the mesophyll;
the analyses remain limited; not all possible relatives palisade parenchyma present or absent; hypoder-
were included. Based on the current information it is mis and epidermis usually well developed and dif-
therefore appropriate to conclude that Phyllocladus is ferentiated. Pollen cones lateral on long shoots or
closely related to Podocarpaceae, but because of the apical on short shoots, often grouped close together
limitations of the molecular studies thusfar under- on long shoots, axillary, solitary or clustered from a
taken and the morphology, the Phyllocladaceae are single bud, catkin-like, deciduous. Microsporophylls
for the present retained as a separate family. The numerous, spirally arranged, (sub)peltate, with 2
other criterion is the evolution of morphological abaxial, free pollen sacs (microsporangia) which
and biological characters. Apart from the phyllo- are longitudinally dehiscent, containing bisaccate
clades (which, being unique within conifers, provide or monosaccate or non-saccate pollen. Seed cones
no information about phylogenetic relationships of woody, small to very large, erect or becoming pendu-
members of that group), there are some quite fun- lous, lateral on long shoots or apical on short shoots,
damental differences that would warrant recogni- often grouped together on long shoots, solitary in
tion at a higher taxonomic rank. These are seen in leaf axils, sometimes long persistent. Bracts spirally
the reproductive morphology (Tomlinson et al., arranged on a woody and integer rachis (breaking
1989), where Phyllocladus has an aril instead of an up in Pseudolarix), remaining small or growing
epimatium, and in the different pollination mecha- with the cone; growing bracts apically differentiated
nisms (Tomlinson et al., 1997). There are a suite of and often exserted. Seed scales axillary to and free
other characters which either unite Phyllocladus from bracts, both persistent or deciduous (Abies,
with Podocarpaceae (Quinn et al., 2002) or sepa- Cedrus); seed scales flattened; apex rounded, more
rate them (Page, 1990), and each author tends to or less acute (Pseudolarix), or differentiated (Pinus).
emphasize those in favour of their respective views Ovules 2 per scale, inverted; seeds slightly flattened,
on the matter. Taxon recognition, despite claims to usually winged (with 1 wing) but in some species of
the contrary, is not a matter to be decided in a purely Pinus effectively wingless due to rudimentary wings
remaining attached to the seed scale. Seedlings with or sometimes epistomatic; seed cones with
varying numbers of cotyledons (224, the highest rudimentary (or small) bract scales and with
number in all plants). Germination epigeal (hypo- rudimentary scales on the pedunculate base,
geal in Keteleeria); young taproot of the seedling pendulous at maturity Picea
with 12 resin canals in the vascular cylinder (stele). 3b. Seeds not separable from relatively small, broad
Diploid number of chromosomes (2n) = 24 (in seed wing; pulvini on shoots weakly developed
Pseudolarix 44 and in Pseudotsuga menziesii 26). or absent; leaves epi/hypostomatic or epi-
amphistomatic with distinct primary stomatal
11 genera: Abies (47 spp.), Cathaya (1 sp.), Cedrus (3 bands; seed cones with small to very large bract
spp.), Keteleeria (3 spp.), Larix (11 spp.), Nothotsuga scales and distinct, leaved peduncles, erect,
52 (1 sp.), Picea (38 spp.), Pinus (113 spp.), Pseudolarix (1 spreading or pendulous at maturity 4
sp.), Pseudotsuga (4 spp.) and Tsuga (9 spp.); total 231 4a. Shoots strongly dimorphic; leaves spirally and
species. remotely arranged on long shoots, in dense
pseudo-whorls on short shoots, deciduous; seed
Distribution cones erect from mostly pendulous branches
Larix
Northern hemisphere (with one equatorial cross- 4b. Shoots monomorphic or weakly dimorphic;
over in N Sumatera). Eurasia & N Africa: Canary leaves spirally arranged, remote or dense, ever-
Islands, Mediterranean coast of N Africa, Europe green; seed cones spreading or pendulous from
(excl. Iceland & Ireland), Turkey, Syria, Lebanon, mostly plagiotropic branches 5
Caucasus, Central Asia, Siberia, Russian Far East 5a. Shoots weakly dimorphic, with (alternate) long
(including Kamchatka), Korea, Japan, Hindu Kush, and short growth; leaves in short growth parts
Himalaya, NE India, China, Taiwan, Indochina, N of shoots in dense tufts; bract scales of seed
Sumatera, Philippines. North America: Canada, cones small, included and simple acuminate
USA (including Alaska, excl. some midwestern Cathaya
states), Mexico. Central America S to Nicaragua; 5b. Shoots monomorphic; leaves equally distant on
Caribbean Islands (excl. Lesser Antilles). all (segments of) shoots; bract scales of cones
large, exserted and trilobate at apex
Key to the genera Pseudotsuga
6a. Mature seed cones pendulous Tsuga
1a. Adult green leaves bundled with 25(-8) 6b. Mature seed cones erect 7
together on dwarf shoots (1 species with a 7a. Pollen cones in (umbellate) clusters from a
single leaf on a dwarf shoot); seed cones bien- single bud; seed cones on mostly long, leaved
nial (rarely triennial), with distinction between peduncles; cone rachis (slowly) disintegrating
each years growth apparent as an umbo and 8
apophysis on each scale; seed held to the wing 7b. Pollen cones solitary; seed cones on short, bare
in a pair of claws Pinus peduncles or sessile; cone rachis persistent 10
1b. Adult green leaves either solitary or in pseudo- 8a. Shoots strongly dimorphic; leaves spirally and
whorls of more than 10 together on short remotely arranged on long shoots, in dense
shoots; seed cones mostly annual (if biennial pseudo-whorls on short shoots, deciduous;
without umbo); seed held to the wing in a cup seed scales deciduous by rapid disintegration of
2 the cone rachis Pseudolarix
2a. Seeds without resin vescicles; seed scales with a 8b. Shoots monomorphic or weakly dimorphic;
broad basis, persistent 3 leaves spirally arranged on long shoots, ever-
2b. Seeds with resin vescicles; seed scales with a green; seed scales (longer) persistent 9
narrow, petiolate basis, persistent or deciduous 9a. Seed cones large, (usually) more than 6cm long
6 and 3 cm wide; leaves with narrowed, twisted
3a. Seeds easily separated from relatively large, base, 24.5mm wide, on weakly developed pul-
narrow seed wing; pulvini on shoots very strongly vini, leaving circular leaf scars after falling
developed; leaves mostly amphistomatic Keteleeria
9b. Seed cones small, (usually) 2.55 1.52.5cm; fleshy or succulent, often brightly coloured recepta-
leaves with oblique, petiolate base, 12 mm cle, or remaining small as part of a small, compound
wide, on more developed pulvini, leaving no cone and either becoming succulent and coloured
distinct leaf scars after falling Nothotsuga (Microcachrys) or leathery and dry (Pherosphaera,
10a. Shoots strongly dimorphic; leaves spirally and Saxegothaea). Seed scales single in the axil of a
remotely arranged on long shoots, in dense bract, bearing a single initially erect but at pollina-
pseudo-whorls on short shoots; seed cones tion more or less inverted ovule, in most genera (not
maturing in two years, disintegrating on tree in Pherosphaera) much expanding after fertiliza-
with deciduous seed scales Cedrus tion to enclose the seed partly or entirely, forming
10b. Shoots monomorphic; leaves spirally arranged the epimatium. Epimatium leathery or fleshy, thin
on long shoots; seed cones maturing in one sea- or very thick, becoming succulent in some genera 53
son, with deciduous bracts and seed scales that do not develop (or only imperfectly develop) a
Abies succulent receptacle, remaining green or variously
coloured. Seeds (except Microcachrys, Pherosphaera
and Saxegothaea) single or sometimes 2, protruding
Podocarpaceae Endl., Syn. Conif.: 203. 1847. (nom. well outside the transformed or reduced cone proper.
cons.). Type: Podocarpus LHr. ex Pers. Seeds proper ovoid, slightly flattened, with a hard,
sclerified seed coat and wingless. Seedlings with
Description 2 cotyledons, germination epigeal. Chromosome
numbers (diploid) 2n = 18 to 38 (but not 28 and 32).
Dioecious or sometimes monoecious, evergreen,
slightly resinous trees or (dwarf) shrubs (one par- 18 genera: Acmopyle (2 spp.), Afrocarpus (5 spp.),
asitic on another member of the family). Bark of Dacrycarpus (9 spp.), Dacrydium (22 spp.), Falcatifolium
trees thin, usually becoming scaly and exfoliating in (6 spp.), Halocarpus (3 spp.), Lagarostrobos (1 sp.),
flakes or strips. Tree trunks monopodial, commonly Lepidothamnus (3 spp.), Manoao (1 sp.), Microcachrys
with orthotropic branching (Rauhs model). Foliage (1 sp.), Nageia (5 spp.), Parasitaxus (1 sp.), Pherosphaera
branches terminating in buds with small, leaf-like (2 spp.), Podocarpus (97 spp.), Prumnopitys (9 spp.),
or more specialized scales (Podocarpus), or termi- Retrophyllum (5 spp.), Saxegothaea (1 sp.) and
nating in reduced leaves. Leaves helically arranged, Sundacarpus (1 sp.); total 174 species.
rarely decussate (Microcachrys), appressed and
imbricate to spreading and remote, shapes highly Distribution
variable, ranging from small, appressed scale leaves,
via thin acicular leaves, to dorsiventrally flattened, Pantropical, extending in the southern hemisphere
linear-lanceolate to broadly lanceolate, large leaves to cool temperate and in the northern hemisphere
(up to 34 9.5cm); venation a single median strand to warm temperate latitudes. Africa: Sub-Saharan
or multiple parallel veins; texture soft and flexible to Africa only, in W Africa from So Thom, S Nigeria
coriaceous and more or less stiff; leaves amphisto- and Cameroon to Angola; in E and S Africa from
matic or hypostomatic. Pollen cones mostly simple, Ethiopia along the Afromontane system to the Cape;
axillary or terminal, solitary or clustered, in some Madagascar. Asia: S India (disjunct in Kerala) and
genera (Nageia, Prumnopitys) forming compound, from the E Himalaya and Assam through S China
racemose units (spikes), sessile or pedunculate, very to Japan; Taiwan, Indochina, Andaman and Nicobar
small to long cylindrical; microsporophylls spirally Islands, and Malesia. Australasia and SW Pacific:
arranged, with 2 abaxial, longitudinally dehiscent Australia (excl. the dry interior and NW), Solomon
pollen sacs; pollen bisaccate or trisaccate (rarely Islands, New Caledonia, Vanuatu, Fiji, Tonga, and
46), or non-saccate (Saxegothaea). Seed cones New Zealand. Americas: From Mexico to Panama,
axillary or terminally, solitary on naked or scaly Caribbean Islands (excl. Bahamas & Turks Caicos
peduncles, or sessile, composed of 2-many alternate Islands); South America along the Andean Cordillera
or helically arranged bracts; fertile bracts 1-many. from Venezuela to S Chili, Venezuelan Highlands,
Sterile and fertile bracts either fusing and forming a SE and SW Brazil.
Key to the genera 11a. Seed cones very small, consisting of multiple
fertile (and some infertile) scales and ovules
1a. Leaves monomorphic, forming a distinct lam- (seeds) 12
ina (blade) wider than thick 2 11b. Seed cones much reduced, usually with only
1b. Leaves monomorphic or dimorphic, at least in one fertile scale producing a single seed 13
part scale-like or more or less acicular, these 12a. Always shrubs, trailing or upright
not or only slightly wider than thick 8 Pherosphaera
2a. Leaves with multiple parallel veins Nageia 12b. Trees (sometimes appearing shrubby with mul-
2b. Leaves with a single (sometimes inconspicu- tiple stems, especially when in cultivation)
ous) midvein 3 Lagarostrobos
54 3a. Seed cones forming an inflated, coloured, suc- 13a. Epimatium forming a yellowish green collar
culent receptacle below the seed(s) when ripe around the erect seed; always trees Manoao
Podocarpus 13b. Epimatium basal, hidden from view by inverted
3b. Seed cones much reduced, not or rarely slightly ripe seed; (dwarf) shrubs or sometimes trees
inflated, or consisting of numerous small, fer- Lepidothamnus
tile scales 4 14a. Leaves acicular, rarely scale-like, never with a
4a. Seed cones consisting of numerous small, fer- distinct lamina Dacrydium
tile scales Saxegothaea 14b. Leaves scale-like and/or with a distinct lamina,
4b. Seed cones much reduced, not or rarely slightly never acicular 15
inflated 5 15a. Epimatium forming a collar around the base of
5a. Pollen cones aggregated in spikes the seeds 17
Prumnopitys 15b. Epimatium surrounding the seeds completely,
5b. Pollen cones in clusters, sessile or very short becoming swollen and succulent 16
pedunculate, or solitary 6 16a. Leaves with well developed lamina always pres-
6a. Leaves in apparent opposite and decussate ent and dominant on all foliage branchlets
pairs due to twisted petioles turning the leaves Acmopyle
in opposite directions Retrophyllum 16b. Laminar leaves present or absent, not domi-
6b. Leaves alternate, not twisted in opposite direc- nant (i.e. scale leaves predominate on some or
tions 7 most foliage branchlets) Dacrycarpus
7a. Leaves with stomata on both surfaces (amphis- 17a. Leaves with well developed lamina always pres-
tomatic Afrocarpus ent and dominant on all foliage branchlets,
7b. Leaves with stomata only on the abaxial (lower) mostly falcate; seed cones forming a swollen
surface (hypostomatic) Sundacarpus receptacle Falcatifolium
8a. Leaves monomorphic (only one type present) 17b. Leaves with well developed lamina absent or
9 present, but not dominant (i.e. scale leaves pre-
8b. Leaves dimorphic (two distinct types present) dominate on some or most foliage branchlets);
14 seed cones much reduced Halocarpus
9a. Small shrubs or dwarf trees with reddish scale
leaves and purple (pen)ultimate branchlets;
seeds terminal, glaucous white Parasitaxus Sciadopityaceae Luerss., Grundz. Bot.: 265. 1877,
9b. Shrubs or trees with green scale leaves on foli- [Sciadopityeae]. Type: Sciadopitys Siebold & Zucc.
age branchlets; seeds lateral or (sub)terminal,
usually not glaucous white 10 Description
10a. Dwarf shrubs, usually creeping, with decussate
scale leaves Microcachrys See the species description.
10b. Shrubs or trees with spirally arranged scale
leaves 11 One genus: Sciadopitys, with 1 species.
Distribution Distribution
As for the species. Western Eurasia & North Africa: Madeira; Atlas Moun-
tains; Europe from Ireland and Portugal to S Scan
dinavia, European Russia and the Caucasus; Turkey,
Taxaceae Gray, Nat. Arr. Brit. Pl. 2: 222, 226. 1822. N Iran. Asia: Hindu Kush, Himalaya, Assam; China,
(nom. cons.). Type: Taxus L. Korea, Russian Far East (Primorye), Japan, Taiwan;
Indochina; Malesia (Sumatera, Sulawesi, Philippines).
Description SW Pacific: New Caledonia. North & Central
America: British Columbia, Washington, Idaho,
Dioecious or rarely monoecious evergreen shrubs W Montana, Oregon, California; E Canada, E USA to 55
or trees, slightly resinous with resin in leaves (with Florida; Mexico, Guatemala, Honduras, El Salvador.
or without resin canals) and arils. Bark thin, exfo-
liating in strips. Branches spreading or ascending, Key to the genera
often with much reiteration; foliage branches termi-
nating in buds with small, imbricate scales. Leaves 1a. Leaves opposite-decussate; pollen cones in
helically or decussately inserted, usually pectinately umbellate clusters of (1-)36 racemes at or just
arranged on plagiotropic branchlets, linear to lan- below the shoot apex Amentotaxus
ceolate, bifacially (dorsiventrally) flattened, with a 1b. Leaves alternate (spirally inserted); pollen
single median vascular bundle (midrib), coriaceous, cones solitary in the axils of leaves along the
hypostomatic, with two separate bands of stomata. shoot (proximally, along the entire length or
Pollen cones lateral on foliage shoots and axillary to distally) 2
foliage leaves, or aggregated in umbellate clusters of 2a. Leaves usually not longer than 3.5cm (rarely to
(1-)36 racemes at or just below the apex of vegeta- 4.5cm); aril surrounding the seed incompletely
tive shoots (Amentotaxus), subtended by small or when mature, leaving the seed well visible 3
large perular scales, short pedunculate, (sub)glo- 2b. Leaves usually longer than 4 cm and up to
bose to short cylindrical. Microsporophylls helically 12(-17) cm; aril surrounding the seed (almost)
or decussately arranged (with secondary displace- completely when mature, leaving the seed (vir-
ment forming 8 ranks), few in number ( but up to tually) invisible 4
60 in Torreya), radially symmetric (peltate) or dor- 3a. Foliage branchlets sub-whorled or subopposite;
siventrally compressed, with 28 pendulous, oblong stomatal bands white; aril of seeds white when
pollen sacs either in radial placement or abaxially, ripe Pseudotaxus
containing spherical, inaperturate pollen. Seed bear- 3b. Foliage branchlets irregularly alternate; sto-
ing structures consisting of axillary dwarf shoots matal bands pale greyish green or pale yellow;
with decussate scales, branching 12 times, usu- aril of seeds red or sometimes orange to yellow
ally strongly reduced, with terminal, erect ovules of when ripe Taxus
which only a single one matures. Ovules at pollina- 4a. Midrib forming a narrow groove on the adaxial
tion time around the base of the integument with a (upper) face of leaves; pollen cones aggregated
ring-like structure. Aril partially or entirely enclos- near the base of new foliage shoots; arils sur-
ing the seed, fleshy or succulent, becoming red, pur- rounding the seeds 1215 79mm, leaving the
ple, or yellow when ripe. Seeds oval to ovoid, slightly seed apex visible Austrotaxus
flattened, with a hard, sclerified seed coat. Seedlings 4b. Midrib on the adaxial face of leaves more or
with 2 cotyledons, germination epigeal. Number less prominently raised; pollen cones form-
of chromosomes (diploid) 2n = 22 (Amentotaxus, ing double rows along the foliage shoots; arils
Torreya) or 24. 1540 1025mm, completely surrounding the
seeds Torreya
Five genera: Amentotaxus (6 spp.), Austrotaxus (1
sp.), Pseudotaxus (1 sp.), Taxus (10 spp.) and Torreya
(6 spp.); total 24 species.
TA XO N OM IC T R E AT M E N T O F G E N E R A A N D SP E C I E S
Abies Mill., Gard. Dict., Abridg. Ed. 4, vol. 1. 1754. Type: Abies alba Mill. [Pinus
picea L.] (Pinaceae).
57
Abies is the classical Latin name for firs. Great Slave Lake to Newfoundland and Appalachian
Mountains in North Carolina. Eurasia and N Africa:
Description from Morocco through S and Central Europe to
Turkey and Lebanon; NE Russia, Siberia and Central
Evergreen monoecious trees with a monopodial, Asia to Sakhalin and Japan; Sino-Himalayan moun-
straight, columnar trunk. Resin canals in bark, leaves, tain system and isolated spots in China, Taiwan.
and seed cones, normally not in wood unless trau-
matic. Branching in rhythmic pseudo-whorls on the Taxonomic notes
trunk (Massarts model), spreading horizontally, with
a dorsiventral symmetry (plagiotropic), branching The genus Abies is the second largest in Pinaceae
again with opposite shoots of lesser potential. (Sub-) (after Pinus). Liu (1971) recognized 39 species,
terminal buds often extremely resinous. Leaves linear, while Rushforth (1987) came to a total of about 55.
more or less flattened, spirally arranged and usually In Farjon (1990) 46 species were recognized. The
twisted at the petiolate base, set on small, circular World Checklist & Bibliography of Conifers (Farjon,
depressions on the shoot which are clearly visible 1998, [2001]) recognized 49, resp. 48 species. Since
after detachment; with two conspicuous white stoma- the monographic work of Liu, several new species
tal bands abaxially and sometimes scattered stomata have been described, mainly from China; further-
also on the adaxial side. Pollen cones usually below more, his treatment of several SW Chinese taxa
seed cones in the tree, axillary, pendulous, solitary, failed to recognize differences on the species level.
catkin-like, with spirally arranged, peltate micro- On the other hand, a number of species mentioned
sporophylls; bearing two pollen sacs with bisaccate by Rushforth have been given infraspecific status
pollen. Seed cones restricted to the upper part of the here, mainly at the subspecies level. Recently, a few
crown, axillary, solitary, sessile or short pedunculate, new species were described from Mexico, but it
erect, disintegrating at maturity when drying. Bracts appears that we have arrived at a phase where these
and seed scales helically attached to a stout or slen- reflect narrower circumscriptions rather than truly
der rachis which remains on the branches. Bracts in new discoveries.
mature cones exserted or sometimes hidden, with a
central cusp; cusp elongated or not. Seed scales cune- Synopsis
ate to reniform, with a pedicellate base. Seeds held
in a membranous cup, covering about 0.7 part of The classification of the genus into sections and
the seed; membrane continued in a well developed, subsections used here to key out the species in the
cuneate-triangular, persistent wing. Seedlings with genus Abies is essentially that of Farjon & Rushforth
(3)48(10) cotyledons. (1989), also used in my book Pinaceae (Farjon,
1990), but with a few emendations. Most notable
46 species of these is the removal of Abies kawakamii from
Taiwan in section Balsamea (subsection Laterales)
Distribution and its tentative placement in section Momi (subsec-
tion Homolepides) as a result of a molecular (nuclear
North America: (disjunct) from Yukon to Arizona DNA) analysis by Xiang et al. (2004). This molecular
and California into Mexico to Honduras; from analysis broadly supported the classification given
by Farjon & Rushforth based on morphological data, Species A. sachalinensis, A. fraseri, A.
but it has demonstrated the anomaly of their place- koreana, A. nephrolepis, A. veitchii
ment of this species. The other changes made here Sect. Grandis Engelm. emend. Farjon & Rushforth
involve a few additions and subtractions of species. Species A. grandis, A. concolor, A.
The classification given below is arranged according durangensis, A. guatemalensis
to hypothetical phylogenetic relationships, but in Sect. Oiamel Franco
the keys to the species the sections and subsections Subsect. Religiosae (Matzenko) Farjon &
are presented in alphabetical order, with sections Rushforth
first and subsections under these. Species A. religiosa, A. vejarii
Subsect. Hickelianae Farjon & Rushforth
58 Genus Abies Mill. Species A. hickelii, A. hidalgensis
Sect. Abies Sect. Nobilis Engelm.
Species A. alba, A. cephalonica, Species A. procera, A. magnifica
A. nordmanniana, A. nebrodensis,
A. cilicica, A. borisii-regis Key to the sections and subsections
Sect. Piceaster Spach emend. Farjon &
Rushforth 1a. Bract cusps more than 2.5cm long. Vegetative
Species A. pinsapo, A. numidica buds fusiform, 12cm long, not resinous
Sect. Bracteata Engelm. emend. Sargent Sect. Bracteata: A. bracteata
Species A. bracteata 1b. Bract cusps much shorter than 1.5 cm.
Sect. Momi Franco Vegetative buds not fusiform, usually much less
Subsect. Homolepides (Franco) Farjon & than 1cm long, often resinous 2
Rushforth, emend. 2a. Seed cones very large (1430 510cm). Leaves
Species A. homolepis, A. kawakamii, on vegetative shoots usually carinate, imbricate
A. recurvata at base, lower leaves curving sideways, upper
Subsect. Firmae (Franco) Farjon & leaves strongly assurgent
Rushforth Sect. Nobilis
Species A. firma, A. beshanzuensis 2b. Seed cones smaller, if longer than 15 cm then
Subsect. Holophyllae Farjon & Rushforth less than 6cm wide. Leaves on vegetative shoots
Species A. holophylla, A. chensiensis, usually flattened 3
A. pindrow, A. ziyuanensis 3a. Seed cones narrowly cylindrical (ratio of length
Sect. Amabilis (Matzenko) Farjon & Rushforth to width greater than 2.5); rachis of cone coni-
Species A. amabilis, A. mariesii cal, slender 4
Sect. Pseudopicea Hickel emend. Farjon & 3b. Seed cones ovoid, conical or broad cylindrical;
Rushforth rachis of cone conical, cylindroconical or fusi-
Subsect. Delavayianae Farjon & Rushforth form, stout 8
Species A. delavayi, A. fabri, A. forrestii, 4a. Seed cones (10)1225(30) cm long, ratio of
A. densa, A. spectabilis, A. fargesii, A. length to width usually 3 or more 5
fanjingshanensis, A. yuanbaoshanensis 4b. Seed cones 312cm long, ratio length to width
Subsect. Squamatae E. Murray usually less than 3 6
Species A. squamata 5a. Bracts exserted and reflexed (usually included
Sect. Balsamea Engelm. emend. Farjon & in A. cilicica), with elongated cusp; cone apex
Rushforth obtuse or acutish (sometimes papillionate)
Subsect. Laterales Patschke emend. Farjon Sect. Abies
& Rushforth, emend. 5b. Bracts always included, cusp short; cone apex
Species A. balsamea, A. lasiocarpa, A. papillionate Sect. Piceaster
sibirica 6a. Seed cones small, (3)48 (1.5)23(4) cm,
Subsect. Medianae Patschke emend. purplish (rarely greenish); bracts yellowbrown.
Farjon & Rushforth Leaves emarginate Sect. Balsamea 7
6b. Seed cones larger, (5)712 35 cm, green, Key to the species of Section Abies
olive green or rarely bluish during the growing
season. Leaves obtuse, acute or emarginate 1a. Bracts included or only slightly exserted. Shoots
Sect. Grandis mostly glabrous; leaves 2.54 cm long, with
7a. Bracts exserted and reflexed; seed scales reni- obtuse or slightly emarginate apex A. cilicica
form Subsect. Medianae 1b. Bracts strongly exserted and reflexed 2
7b. Bracts included; seed scales cuneateflabellate 2a. Leaves 12.2 cm long, 23.5 mm wide, rigid;
Subsect. Laterales apex acute or obtuse, not emarginate. Seed
8a. Seed cone rachis thick, fusiform or cylin- cones small, 712 34cm A. nebrodensis
droconical; seed scales usually apically thick- 2b. Leaves (usually) longer than 2cm, up to 2.5mm
ened. Shoots usually stout wide, acute to emarginate. Seed cones usually 59
Sect. Pseudopicea 9 larger 3
8b. Seed cone rachis less thick, conical; seed scales 3a. Seed cones long and narrow (ratio length to
thickest at or below the middle. Shoots usually width 34), width 35 cm. Leaves acute or
thin 10 emarginate 4
9a. Bark exfoliating in large papery flakes Subsect. 3b. Seed cones large, (10)1220 46 cm (ratio
Squamatae: A. squamata length to width up to 3 at most). Leaves emar-
9b. Bark different Subsect. Delavayianae ginate, on shaded, vegetative shoots pressed
10a. Bracts usually exserted with a broad cusp (but forward, covering them A. nordmanniana
not in A. vejarii subsp. mexicana). Branchlets 4a. Leaves on shaded shoots emarginate, mostly in
purplish brown Sect. Oiamel 11 pectinate arrangement with overlapping ranks.
10b. Bracts included with a small cusp (which may Young shoots pubescent 5
be slightly exserted) or if exserted, cone 4b. Leaves acute, rarely obtuse, more radially
oblongconical and green 12 spreading. Young shoots glabrous A.
11a. Leaves emarginate or obtuse; resin canals in the cephalonica
leaves 410(12) Subsect. Hickelianae 5a. Leaves on the adaxial (upper) side lustrous
11b. Leaves acute; resin canals in the leaves 2 green, without any stomata; apex of leaves on
Subsect. Religiosae non-coning shoots slightly emarginate A. alba
12a. Young shoots uniformly pubescent; leaves 5b. Leaves on the adaxial side with a few stomata;
densely crowded above the shoot, directed for- apex of all leaves entire A. borisii-regis
ward Sect. Amabilis
12b. Young shoots glabrous or weakly pubescent in
Key to the species of Section Amabilis
grooves; leaves not dense, more pectinate
Sect. Momi 13 1a. Cones large, 915 58cm. Leaves 23cm long;
13a. Seed cones oblongconical; bracts strongly buds ca. 5 4mm A. amabilis
exserted Subsect. Firmae 1b. Cones smaller, 49 24.5cm. Leaves shorter
13b. Seed cones ovoidoblong to cylindrical; bracts (usually 12 cm long); buds small, 23 mm
included or cusps just exserted near the base of long A. mariesii
the cone 14
14a. Leaves 1.53(3.5) cm long. Seed cones vio-
Key to the species of Section Balsamea,
letblue or purple, oblong cylindrical to
Subsection Laterales
ovoidoblong, 2.53.5(4) cm wide
Subsect. Homolepides 1a. Leaves on vegetative shoots mostly pectinate, a
14b. Leaves 25(9) cm long. Seed cones yellowish few above the shoot directed forward, short
green to violetblue, cylindrical or ovoid and stiff. Seed cones (2.5)58 23cm
cylindrical, 3.56cm wide A. balsamea
Subsect. Holophyllae 1b. Leaves on vegetative shoots mostly assurgent,
short or longer, flexible. Seed cones mostly
larger 2
2a. Leaves strongly assurgent, densely covering the 3a. Leaves green or glaucous green, stomata on
shoot; buds usually larger than 3mm both sides, leaf apex entire A. concolor
A. lasiocarpa 3b. Leaves dark glossy green above, stomata on one
2b. Leaves slightly assurgent, the lower leaves side only, leaf apex emarginate A. grandis
spreading laterally, more remote, leaving the
shoot visible; buds 23mm long A. sibirica Key to the species of Section Momi,
Subsection Firmae
Key to the species of Section Balsamea,
Subsection Medianae 1a. Bracts oblanceolate, slightly exserted and not
reflexed. Buds large (max. 10 5mm). Leaves
60 1a. Seed cones with large, strongly exserted, light green A. firma
reflexed and light coloured bracts 2 5b. Bracts spathulate, exserted and reflexed at the
1b. Seed cones with smaller, less exserted and upper margin. Buds smaller. Leaves dark green
reflexed, usually darker coloured bracts 3 A. beshanzuensis
2a. Seed cones very numerous, even on the lower
branches, often crowded together, appearing Key to the species of Subsection Holophyllae
soon on young trees; bracts leaving the apex of
the seed scales mostly free. Leaves on vegetative 1a. Leaves on vegetative shoots entire 2
shoots emarginate A. koreana 1b. Leaves on vegetative shoots emarginate or
2b. Seed cones absent from the lower branches, less bifid 3
numerous; bracts very broad, covering most of 2a. Seed cones oblongcylindrical, 34.5cm wide.
the seed scales. Leaves on vegetative shoots Leaves on leading and coning shoots strongly
mostly obtuse A. fraseri assurgent, acute or mucronate A. holophylla
3a. Young shoots redbrown or brown, rarely 2b. Seed cones broader, 56(7) cm. Leaves slightly
greybrown. Leaves often strongly twisted at assurgent and bifid, emarginate or sometimes
base; cones usually ellipsoidcylindrical. Bracts obtuse A. pindrow
variously exserted A. sachalinensis 3a. Leaves 33.5mm wide, 24.8cm long, obtuse or
3b. Young shoots yellowish or greenish brown. rarely slightly emarginate. Ripe seed cones dark
Leaves less twisted; cones cylindrical. Bracts brown; bracts spathulate, with 910 mm wide
slightly exserted 4 apex A. ziyuanensis
4a. Young shoots densely yellowish pubescent. 3b. Leaves 2.53mm wide, when wider, then longer
Leaves dark green above. Bracts slightly than 4.5cm, at least on vegetative shoots emar-
exserted A. veitchii ginate. Ripe seed cones (light) cinna-
4b. Young shoots minutely pubescent in grooves. monbrown; bracts not spathulate
Leaves light green above. Bracts entirely A. chensiensis
included or only the cusps of the bracts
exserted A. nephrolepis Key to the species of Subsection Homolepides
Key to the species of Section Grandis 1a. Leaves spreading pectinately or at least parted
above the shoot A. homolepis
1a. Leaves very thin, less than 2 mm wide, very 1b. Leaves crowded and curved upwards or some-
flexible 2 times reflexed above the shoot 2
1b. Leaves thicker, 23mm wide, more rigid 3 2a. Leaves on vegetative shoots 1.52 mm wide,
2a. Young shoots (light) greenish to reddish brown. curved upwards A. kawakamii
Leaves 1.22 mm wide, emarginate or obtuse. 2b. Leaves on vegetative shoots 1.92.5 mm wide,
Bracts slightly exserted A. guatemalensis recurved or reflexed A. recurvata
2b. Young shoots dark purplish red or red-brown.
Leaves 11.6mm wide, slightly acute or obtuse.
Bracts included A. durangensis
Key to the species of Section Nobilis more radially, shorter or with strongly revolute
margins 2
1a. Bark of old trees (in their native habitat!) grey- 2a. Leaves with (strongly) revolute margins; sto-
ish brown. Bracts of seed cones far exserted and matal bands (partly) hidden, niveous white 3
strongly reflexed, covering much of the seed 2b. Leaves not revolute or with only very slightly
scales. Leaves partly grooved above A. procera recurved leaf margins, stomatal bands entirely
1b. Bark of old trees (dark) redbrown. Bracts of visible, white or greyish white 4
seed cones included or slightly exserted and 3a. Bracts abruptly ending in an elongated, awllike
reflexed. Leaves not grooved A. magnifica cusp, usually exserted (at least with the long
cusp). Young shoots dark redbrown
Key to the species of Section Oiamel, A. delavayi 61
Subsection Hickelianae 3b. Bracts gradually ending in a short cusp, not or
slightly exserted. Young shoots yellowish
1a. Bracts of seed cones exserted A. hickelii brown A. densa
1b. Bracts of seed cones included A. hidalgensis 4a. Seed cones green or yellowish green in the
growing season; bracts exserted and reflexed
Key to the species of Section Oiamel, A. yuanbaoshanensis
Subsection Religiosae 4b. Seed cones purple or purplish blue in the grow-
ing season; bracts exserted or sometimes
1a. Seed cones (8)1016 46cm; bracts strongly included, not reflexed 5
reflexed, partly covering the seed scales. Leaves 5a. Young shoots usually purplish, reddish, or dark
pectinately arranged, usually 1.53 cm long, orangebrown 6
leaving the shoot visible from above 5b. Young shoots yellowish brown, mostly glabrous
A. religiosa A. fabri
1b. Seed cones 612(15) 46(7?) cm; bracts not 6a. Seed cones small, 59 34 cm; bracts with
reflexed. Leaves spreading but not pectinate, rounded or emarginate apex ending in a small,
usually 12cm long, covering the shoot above abrupt cusp 7
A. vejarii 6b. Seed cones 610(14) 45(6) cm, or if less
than 8 4cm, with differently shaped bracts
Key to the species of Section Piceaster A. forrestii
7a. Leaves 12.5cm long, rarely up to 4.3cm, very
1a. Leaves carinate, obtuse or slightly acute, densely set in overlapping, pectinate rows.
amphistomatic,very rigid and radial. Buds res- Bracts mostly included or slightly exserted,
inous. Seed cones 914 34cm A. pinsapo with very short cusps A. fanjingshanensis
1b. Leaves flattened, weakly amphistomatic, on 7b. Leaves (1)1.53(4.5) cm long, especially on
shaded shoots less radial. Buds not resinous. shaded shoots, less dense. Bracts exserted, with
Seed cones usually larger, 1218 46cm much longer cusps A. fargesii
A. numidica
Key to the species of Section Pseudopicea, Abies alba Mill., Gard. Dict., ed. 8: Abies No. 1.
Subsection Delavayianae 1768. Type: Habitat in Alpinus Helvetiae, Sueviae,
Bavariae, Scothiae, Herb. Clifford 449 Abies 2
1a. Seed cones large, usually 1017 47cm; bracts (lectotype BM). Fig. 1
included. Leaves pectinately arranged, 2.56cm
long, 2.23.5 mm wide, bifid or emarginate,
with slightly recurved margins Abies alba Mill. subsp. apennina Brullo, Scelsi &
A. spectabilis Spampinato, La Vegetazione dell Aspromonte: 41.
1b. Seed cones smaller, if larger than 10 5cm usu- 2001.
ally with exserted bracts. Leaves spreading
Etymology slightly recurved at maturity. Seeds cuneate, angular,
79mm long, yellowish brown; seed wings cuneate,
The species epithet means white and perhaps refers 1015mm long, reddish yellow.
to the light grey bark.
Distribution
Vernacular names
Europe: From the Pyrenees to the Carpathians, Italy
Silver fir; Sapin pectin (French); Weisstanne (Apennines) and in the Balkan Peninsula to Bulgaria
(German) and N Greece.
TDWG codes: 11 AUT-AU AUT-LI CZE-CZ CZE-SL
62 Description GER HUN POL SWI 12 COR FRA-FR SPA-AN SPA-SP
13 ALB BUL GRC ITA-IT ROM YUG-BH YUG-CR
Trees to 55(60?) m tall, d.b.h. to 22.5 m; trunk YUG-KO YUG-MA YUG-MN YUG-SE YUG-SL 14
monopodial, straight, columnar, terete; crown UKR-UK
conical, but flat topped (storks nest) in most old
trees. Bark in young trees greyish, smooth, in old Ecology
trees greyish brown, light or dark, becoming scaly
and fissured below. Branches of first order hori- Primarily a montane species, occurring between 500
zontal, descending, ascending towards the top of and 1500 m a.s.l., but as low as 300 m in the Bavarian
the tree; branches of second order slender, spread- Forest and up to 1950 m in the Pyrenees. Soils are
ing horizontally. Branchlets slender, flexible, stout usually well drained sandy loams from silicate rocks
and firm on cone bearing branches, yellowish grey or limestone derivatives. The climate is cool tem-
or grey, nearly smooth, striated, with short, yellow- perate, comparatively humid (precipitation often
ish or brown pubescence or occasionally glabrous; >1000mm/year), with abundant snowfall but mod-
leaf scars broadly elliptic or circular. Vegetative erately low temperatures in the winter. The species
buds ovoid or conical, ca. 6 4mm, not resinous or often forms large forests, either pure, mixed with
slightly resinous especially on cone bearing branches; other conifers (Picea abies, locally Pinus sylvestris)
bud scales broadly ovate, obtuse, brown, persisting or mixed with broad-leaved trees (Fagus sylvatica),
several years. Leaves spirally arranged, on vegeta- in a belt between deciduous forest in the valleys
tive branches pectinate in two lateral sets, on leading and coniferous forest composed of other species of
shoots radial, on coning shoots radial and assurgent, Pinaceae towards the tree limit. Of the European
(1.2)1.53(3.5) cm long, 1.52.6mm wide, (slightly) native conifers, Abies alba is most capable of com-
twisted at base, linear, flattened, grooved and lus- peting with Beech (F. sylvatica) at altitudes where
trous dark green on the adaxial (upper) face, whit- the latter becomes less vigorous and is the first of the
ish green below; apex slightly emarginate (obtuse conifers to appear among them (Ellenberg, 1988).
or acute on coning shoots). Stomata in two bands
separated by a midrib on the abaxial leaf face. Pollen Conservation
cones lateral, crowded, 2cm long, greenish yellow
with purple red microsporophylls. Seed cones lat- IUCN: LC
eral, erect; peduncles short, slightly curved, cylin-
drical, with conical or obtuse apex, 1015(20) cm Uses
long, 35(6) cm wide, yellowish green when imma-
ture, sometimes with purple tinge, ripening to light White fir is an important timber tree in western and
brown or reddish brown; cone rachis persistent, nar- central Europe, where most forests are semi-natural
rowly conical, dark brown. Seed scales cyathiform, and managed with a view to encourage fir regenera-
the upper ones more cuneate, length width at mid- tion and growth at the expense of competitors. It is
cone 2.53 2.53.5cm; surface smooth, pulverulent not very successful as a plantation forestry tree out-
especially on the exposed part; upper margin entire; side its natural areas of occurrence; one cause of this
base long pedicellate. Bracts linear-spathulate, with may be damage done by insect pests, which may be
long, caudate cusp, 2.53.5(4) cm long, exserted, more prolific in monocultures and in areas with mild
winters such as in the British Isles. Its most famous ones pectinate, the upper ones directed forward, cov-
use in the past was for the masts of 17th century ering the branches, on coning shoots radial, curved,
ships. Most of its wood today is used for plywood the upper ones assurgent, 23cm long, 1.52mm
and veneer as it is evenly grained, light, and easily wide, twisted at base, linear, flattened, grooved and
worked. Minor uses are for soundboards in musical dark lustrous green above, two silvery white lines
instruments, boxes, wood carving, and sometimes below; apex slightly notched or obtuse. Stomata
for joinery. Distillation of the leaf essential oils has in in two bands separated by a midrib on the abaxial
the past been used to produce remedies for sprains (lower) leaf face, none or a few on the adaxial side.
and bruises. In Britain it was used as a Christmas Pollen cones lateral in leaf axils mostly solitary but
tree in the 19th century when Prince Albert popular- numerous, 11.5cm long, with red microsporophylls.
ized the tradition; its use has declined since in favour Seed cones lateral, erect, mostly towards the ends of 63
of the more quickly and cheaply produced Norway branches on short peduncles, ovoid-cylindrical or
spruce. In horticulture White fir is uncommon and oblong, with obtuse or papilliform apex, 915cm
mostly restricted to arboreta; a number of cultivars long, 58cm wide; purple or rarely greenish when
are known but few are common in the trade. immature, purplish brown when maturing, turning
brown when ripe; cone rachis persistent, cylindric
conical, dark or light brown. Seed scales cyathiform-
Abies amabilis Douglas ex J. Forbes, Pinetum flabellate or cuneate, recurved when ripe, length
Woburn.: 125, t. 44. 1839. Picea amabilis Douglas width at mid-cone 22.8 2.53.5cm; surface
ex Loudon, Arbor. Frut. Brit. 4: 2342. 1838. smooth, often densely covered with resin, puberu-
Type: A collection made by David Douglas, not lous on both surfaces; upper margin entire, strongly
designated. Fig. 2 incurved; base long pedicellate. Bracts obovate-
oblong, 11.5cm long, included, with very short
Etymology cusps. Seeds oblong, 1012mm long, light brown;
seed wings quadrangular, broad, 18 20mm, yellow
The species epithet means lovely and was coined by or pale brown.
its discoverer David Douglas.
Distribution
Vernacular names
Pacific coast region of W North America, from
Pacific silver fir, Lovely fir, Cascades fir extreme SE Alaska to N California.
TDWG codes: 70 ASK 71 BRC 73 ORE WAS 76 CAL
Description
Ecology
Trees to 4580 m tall, d.b.h. to 23 m; trunk monopo-
dial, straight, columnar, massive, terete; crown nar- Abies amabilis occurs from sea level near the coast
rowly conical or irregular in old trees. Bark in young to 330 m a.s.l. in SE Alaska, in Oregon from 250 m
trees smooth, ash grey or whitish, with resin blisters, to 1830 m a.s.l. on the western slopes of the Cascade
in old trees deeply furrowed, brown, divided into Range. It grows on different mountain soils, usually
small plates. Branches of first order horizontal, rela- of glacial origin and acidic. The climate is extremely
tively short, thick, curved downward on lower part wet maritime, with 1500 to 4000mm annual pre-
of the trunk; branches of second order short, curved, cipitation, much of it as snow. It is a constituent of
spreading. Branchlets slender, light greenish brown the mixed coniferous forests with among other coni-
or greyish, changing to reddish brown, ridged and fer tree species Tsuga heterophylla, Picea sitchensis,
grooved, remotely or more densely pubescent with Pseudotsuga menziesii, Thuja plicata, Chamaecyparis
light hairs; leaf scars circular, reddish. Vegetative nootkatensis, Abies grandis, A. magnifica; and with
buds subglobose, ca. 5 4mm, very resinous; bud A. lasiocarpa and Tsuga mertensiana at higher ele-
scales dark purple, persisting several years. Leaves vations, but unlike the latter two not reaching the
spirally arranged, on vegetative branches the lower tree line.
Conservation 2mm wide, twisted at base, linear, flattened, dark
green above, two white bands separated by green
IUCN: LC midrib below; apex obtuse or emarginate (acute or
acuminate on cone bearing branches). Stomata on
Uses the adaxial (upper) surface scattered along a medial
groove, increasing towards apex, on lower surface in
In the timber industry no distinction is made two bands separated by a midrib. Pollen cones lat-
between this species and Western hemlock as both eral, in leaf axils, pendant, crowded and numerous,
conifers have similar wood properties. This wood is 0.6cm long, yellow, with purple microsporophylls.
in use for various construction applications such as Seed cones lateral, erect; peduncles very short; shape
64 plywood, veneer, sub-flooring and sheathing. It con- cylindrical, narrowing towards an obtuse apex, (2.5
tains little or no resin and is light in colour and easily )58cm long, (1.5)23cm wide, green, tinged with
worked. Together with Western hemlock, substan- purple, or violet-purple when immature, ripening to
tial quantities of wood go to the kraft pulp industry. (purplish) brown; cone rachis persistent, narrowly
As an ornamental tree it is uncommon, performing conical, blackish brown. Seed scales cuneate, flabel-
only in cool and wet maritime climate such as pre- late or rhombic-orbicular, length width at mid-
vails in the west of Scotland. cone 1.21.4 1.41.7cm; surface smooth, striate on
lower part, pubescent with brown hairs on exposed
part; upper margin entire, strongly curved inward;
Abies balsamea (L.) Mill., Gard. Dict., ed. 8: Abies base pedicellate. Bracts broad ligulate, with a short
No. 3. 1768. cusp, 0.61cm long, entirely included or sometimes
exserted and recurved. Seeds cuneate, 34 long, dark
Etymology grey; seed wings cuneate, oblique at apex, 710mm
long, greyish brown.
The species epithet refers to the balsam i.e. its corti-
cal resin which has medicinal properties. Distribution
Balsam fir is an economically important conifer. Its Abies balsamea (L.) Mill. var. phanerolepis
wood, although of modest size, is used in light-frame Fernald, Rhodora 11: 201. 1909. Abies intermedia
construction and for pulpwood. It is also popular as Fulling, Castanea 1: 93. 1936; Abies balsamea (L.) 65
a Christmas tree and is one of the top three species Mill. f. phanerolepis (Fernald) Rehd., Man. Cult.
grown for this purpose in E North America. The fra- Trees, ed. 2: 16. 1940; Abies phanerolepis (Fernald)
grant needles are partly responsible for this popular- T. S. Liu, Monogr. Gen. Abies: 316. 1971; Abies
ity, they are also used to stuff pillows sold as souvenirs balsamea (L.) Mill. subsp. phanerolepis (Fernald)
in New England. Canada balsam, the aromatic and E. Murray, Kalmia 12: 18. 1982. Type: Canada,
soft terpenoid resin collected from blisters in the Quebec, Perc Mt., M. L. Fernald & J. F. Collins 860
bark, is especially important in Quebec. Its medici- (holotype GH).
nal properties were known to Native Americans,
who used it as an antiseptic wound dressing as Description
well as internally for various ills. Lewis and Clark
had it in their medicine box on their famous over- Seed cones 25.5cm long, 1.52cm wide, violet-pur-
land expedition to the Pacific Ocean in 180405. In ple when young, greyish or blackish brown at matu-
modern Western society its medicinal use has been rity, brown pubescent; bracts exserting, with long,
replaced by other salves; the resin is now used to seal reflexed cusps.
microscopic glass slides with biological preparates.
In horticulture, Balsam fir is less valued; this fir is Taxonomic notes
apparently short lived when planted in gardens and
only a few dwarf cultivars are known. This fir has been interpreted by Liu (1971) as a hybrid
between A. balsamea and A. fraseri; this taxonomy
2 varieties are recognized: was accepted in my book Pinaceae (Farjon, 1990)
but is here rejected in favour of an infraspecific
taxon under A. balsamea, with which it shares more
Abies balsamea (L.) Mill. var. balsamea. Pinus character states. Only the smaller cones with exsert-
balsamea L., Sp. Pl. 2: 1002. 1753. Type: Habitat in ing but small bracts remind of A. fraseri and a hybrid
Virginia, Canada, leg. ign. LINN 1135.14 (lectotype origin of this variety has not been demonstrated
LINN). genetically.
Description Distribution
Seed cones 58cm long, 23cm wide, green with Canada: Quebec, Perc Mt., Mont Blanc (Matane
a purplish tinge when young, brown at maturity; Co.); USA: Virginia, West Virginia.
bracts included or only the cusps exserting. TDWG codes: 72 QUE 75 WVA 78 VRG
Distribution Conservation
The epithet refers to the name of the mountain on This species resembles A. firma in several charac-
which it was found. ters. Wu (1976) related this species to A. fabri on
66 the grounds of its recurved cone bracts, but both
Vernacular names colour and shape of the cones are quite different and
resemble the cones of A. firma, as do the broad, flat
Baishan fir; Baishanzu lengshan (Chinese) and emarginate leaves. Hence, Farjon & Rushforth
(1989) classified it in the section Momi, subsection
Description Firmae.
plate 1. Abies bracteata. 1. Habit of tree. 2. Foliage branch. 3. Seed cone; seed scales with bracts.
4.Detail of buds and leaves. 5. Pollen cone. 6. Seeds. 7. Leaves.
Distribution later in the 20th century when renewed seed collect-
ing was undertaken. Trees in cultivation often grow
USA: California (Santa Lucia Mts.). much faster and taller than in their natural habitats
TDWG codes: 76 CAL in the Santa Lucia Mountains.
Ecology
Abies cephalonica Loudon, Gard. Mag. & Reg.
This fir species occurs in the Upper Sonoran and Rural Domest. Improv. 14: 81. 1838 [Arbor. Frutic.
Transition Life Zones of the coastal Santa Lucia Brit.]. Abies alba Mill. var. cephalonica (Loudon)
Mountains, in canyon bottoms and on rocky slopes Richt., Pl. Europ. 1: 5. 1890. Type: not designated.
70 at elevations between (180)600 and 1570 m a.s.l. It Fig. 4
grows on rocky, clay or loam rich mountain soils,
which can be very dry in summer. It usually grows Etymology
within reach of ground or seepage water or near
intermittent canyon bottom streams. The climate is The species epithet denotes its locus classicus, the
warm, with dry summers and moderate, moist win- island of Cephalonia in Greece.
ters, the precipitation ranges from 500 to 1300mm
annually. It is a constituent of a mixed forest with Vernacular names
Pinus coulteri, P. ponderosa, Pseudotsuga menziesii,
Quercus spp., Lithocarpus densiflorus, Acer macro- Greek fir, Grecian fir; Kukunaria (Greek)
phyllum, and other species.
Description
Conservation
Trees to 3035 m tall, d.b.h. to 1.52 m; trunk mono-
Abies bracteata has a limited range, consisting of a podial, straight, columnar, terete; crown pyramidal,
reasonably large number of small and often quite old trees flat topped. Bark of young trees greyish,
disjunct subpopulations, each containing from a few tinged with pink or brown, smooth, finally in old
score to several hundred trees. No commercial exploi- trees fissured below. Branches of first order long,
tation threatens this species and most populations are spreading horizontally, pendant below; branches of
within protected areas or on public lands where the second order spreading or pendant. Branchlets stout,
management is aimed at preserving these remarkable shaded ones slender, shining pale brown or reddish
trees. However, they are susceptible to forest fires, brown, rarely yellowish, prominently grooved, gla-
which in much of central and southern California brous; leaf scars circular. Vegetative buds ovoid or
have increased in intensity and extent due to a com- conical, ca. 6 45mm, with yellowish resin; bud
plex of largely anthropogenic factors. A hot forest fire scales ovate, acute, reddish brown and persistent
at the wrong time in the wrong location could wipe for several years. Leaves spirally arranged, spread-
out an entire subpopulation. However, it appears that ing radially, more or less pectinate below, assurgent
nearly all stands are in situations where a fire would and curved on coning shoots, (1.5)23(3.5) cm
not reach or have limited destructive power. long, 22.5mm wide, twisted or curved at base, lin-
IUCN: NT ear, curved and flattened, shining dark green above,
with two whitish green stomatal bands separated by
Uses a midrib below; apex acute, rarely obtuse. Stomata
few on the adaxial (upper) surface near the apex,
Santa Lucia fir is not used for timber but it is an attrac- two bands separated by a midrib below. Pollen cones
tive and unusual species much valued in collections lateral, crowded on the underside of branches, 1.2
for botanic gardens and arboreta. It was successfully 1.8cm long, yellow with red microsporophylls. Seed
introduced and raised by the famous tree nursery cones lateral, erect; peduncles short; cone shape
of Veitch & Son near Chelsea in England in 1853, cylindrical, 1016(20?) cm long, 3.55cm wide;
but it only became more common in horticulture apex obtuse or papilliform, greenish brown when
immature, maturing to light brown; cone rachis per- nigra, but higher in the mountains it usually forms
sistent, narrowly conical, brown; seed scales cuneate pure forests with occasional Juniperus oxycedrus.
near the top, cyathiform at mid-cone, length width
at mid-cone 2.53.5 23cm; surface smooth, with Conservation
yellowish brown pubescence on the exposed part;
upper margin undulate or entire, convex; base short Abies cephalonica has a limited distribution, but its
pedicellate. Bracts linear-spathulate, with long cusp, actual extent of occurrence (EOO) is difficult to
34cm long, exserted and recurved. Seeds narrowly establish due to inferred past and perhaps present
cuneate, 8 5mm, brown; seed wings cuneate or hybridization with A. alba. Where it is considered
oblong, 1520 810mm, lustrous light brown. genetically pure, the introduction of firs from other
sources would potentially threaten this species; it 71
Taxonomic notes is therefore imperative for foresters to refrain from
such introductions among or near such populations.
The genetic purity of A. cephalonica is apparently The situation becomes more obscure in the northern
lacking in much of the presumed former range of Peloponnisos and in Sterea Hellas where it is found
this species (Mitsopoulos & Panetsos, 1987). There that both A. cephalonica and A. borisii-regis already
appears to be a north-south cline of character states, occur. Climate change may influence this situation,
reflecting introgression with A. alba genes with with drier conditions presumably benefiting A. ceph-
decreasing influence on morphological traits. Pure alonica and wetter conditions A. alba, and therefore
A. cephalonica populations occur on Cephalonia ongoing introgression of the genes of A. alba into the
(the locus classicus), Euboea and in the far south of remaining populations of A. cephalonica.
Peloponnesos, but individual trees with genuine char- IUCN: LC
acters of this species occur mixed with intermediate
forms well into Macedonia. Variation is considerable Uses
in much of the area where these hybrid forms are sup-
posed to occur. This hybrid has been formally named Greek fir, due to its rarity, is of limited commercial
as Abies borisii-regis Mattf. and was originally value as a timber tree. Crosses with other European
described from the Rhodope Mountains in Bulgaria. firs (A. nordmanniana, A. pinsapo) have been estab-
Other botanists described the hybrid as a variety or lished and tried or used in plantation forestry as well
subspecies of A. alba (A. alba var. acutifolia Turrill). as the natural hybrid A. borisii-regis. The latter is
more widespread and, together with European fir (A.
Distribution alba), exploited for timber in its natural range. Abies
cephalonica was widely planted as an ornamental
Greece (Cephalonia, Euboea, Sterea Hellas, tree in Europe during the 19th century, distributed
Peloponnisos). from seed collected on Cephalonia. Such trees are
TDWG codes: 13 GRC now becoming rare because seedlings from them
will not be genetically pure and new collections are
Ecology seldom made. A few cultivars are known but these,
too, are rare.
Abies cephalonica is a montane species, occurring
from (600)8002000(2100) m a.s.l. in the high
mountains of Greece. Soils are usually well drained Abies chensiensis Tiegh., Bull. Soc. Bot. France 38:
and calcareous, but in the north, where introgres- 413. 1892.
sion with A. alba seems to have occurred, also on
siliceous soils, which may be slightly acid. The cli- Etymology
mate has relatively dry summers and wet winters,
with annual precipitation from 700 to 1500mm. At The species epithet refers to the Chinese province
lower elevations A. cephalonica mixes with Fagus ori- (now spelled Shaanxi) from where this species was
entalis, Quercus spp., Castanea sativa, and also Pinus first described.
Vernacular names Ecology
Shensi fir; qin ling leng shan (Chinese) This species occurs in high mountain ranges of the
SW Plateau of China between 2100 and 3000 m a.s.l.,
Description on grey-brown mountain podzols, brown earth or
lithosols. The climate is cold and moist, with annual
Trees to 50(70?) m tall, d.b.h. to 22.5 m; trunk precipitation between 1000 and 2000mm. It is a rare
monopodial, straight, columnar, terete; crown pyra- species, usually mixed with Picea spp., Abies fargesii
midal, becoming flat topped in old trees. Bark of var. sutchuenensis, Tsuga chinensis, Larix potaninii at
young trees dark grey, smooth, fissured in old trees. high elevations, and Betula spp. at lower elevations;
72 Branches of first order short, massive; branches of also as a pure forest in Tsin-ling Shan (Wang, 1961).
second order stout, spreading. Branchlets stout,
firm, yellowish or grey, shining, ridged between Uses
leaf scars, glabrous or slightly pubescent at first; leaf
scars roundish with a light centre. Vegetative buds This fir has been logged heavily for timber in north-
conical or ovoid, 10 6mm or larger on some lead- ern China. As of most tall firs, the wood is soft,
ing shoots, (slightly) resinous; bud scales triangu- even-grained and suitable for many applications in
lar or broadly ovate, brown or reddish, persisting construction, veneer, panelling etc. The species is
several years. Leaves spirally arranged, the lower uncommon in cultivation outside China and mostly
ranks pectinate at right angles to shoot, the upper restricted to botanic gardens and arboreta.
leaves shorter and directed forward, assurgent or
nearly erect on coning shoots, (1.3)24.5(7.5) cm Three subspecies are recognized:
long, 2.53mm wide, twisted at base, ligulate linear,
flattened, grooved and shining green above, whit- Abies chensiensis Tiegh. subsp. chensiensis. Type:
ish green below, emarginate or obtuse, occasion- China, Shaanxi, Qinling Shandi, Abb David 918
ally acute (coning shoots). Stomata in two broad (holotype P).
bands divided by a midrib below. Pollen cones lat-
eral, crowded, 1cm long. Seed cones lateral, erect on Description
short peduncles, oblong-ovoid or cylindrical, with
a truncate apex, (7)811(14) cm long, (3)45cm Leaves (1.3)24.5(5) cm long; resin canals in leaves
wide, green when immature, ripening to cinnamon of coning shoots medial. Seed cones (7)811(12)
brown; cone rachis persistent, cylindric-conical, cm long.
dark brown. Seed scales broadly ovate-cuneate to
cyathiform, length width at mid-cone 2.53 Distribution
33.5cm; surface smooth, tomentose-puberulent;
upper margin rounded, obscurely serrate; base pedi- China: SE Gansu, Henan (Nexiang), W Hubei, S
cellate. Bracts short, 2-laminated, with short cusp, Shaanxi, W Sichuan.
length 11.5cm, entirely included. Seeds obovoid, 10 TDWG codes: 36 CHC-HU CHC-SC CHN-GS
5mm, brown; seed wings obdeltoid, 20 10mm, CHN-SA CHS-HE
pale brown.
Conservation
Distribution
Logging and deforestation have depleted this most
China: SE Gansu, Henan, W Hubei, Shaanxi, W common and widespread subspecies, but to an
Sichuan, NW Yunnan, SE Xizang [Tibet]; NE India: unknown extent. A logging ban imposed by the cen-
Arunachal Pradesh. tral government applies to all native forest with this
TDWG codes: 36 CHC-HU CHC-SC CHC-YN and other conifer species.
CHN-GS CHN-SA CHS-HE CHT 40 EHM-AP IUCN: LC
Abies chensiensis Tiegh. subsp. salouenensis Taxonomic notes
(Bordres & Gaussen) Rushforth, Notes Roy. Bot.
Gard. Edinburgh 41: 539. 1984. Abies salouenensis Rushforth (1984) described this subspecies from the
Bordres & Gaussen, Trav. Lab. Forest. Toulouse T. main massif of the Lijiang Shan (Yulongxue Shan,
1 (4, 15): 4. 1947; Abies ernestii Rehd. var. salouenen- 5596 m) in NW Yunnan. It has somewhat larger cones
sis (Bordres & Gaussen) W. C. Cheng & than the type subspecies and marginal resin canals in
L. K. Fu, Fl. Reipubl. Pop. Sin. 7: 93. 1978; Abies the leaves of cone-bearing shoots. In Flora of China 4
recurvata Mast. var. salouenensis (Bordres & (1999) where no subspecies of A. chensiensis are rec-
Gaussen) C. T. Kuan, Fl. Sichuan. 2: 48. 1983; Abies ognized, this taxon has been instead treated under
chensiensis Tiegh. var. salouenensis (Bordres & Abies ernestii Rehd. var. salouenensis (Bordres &
Gaussen) Silba, Phytologia 68: 10. 1990. Type: Gaussen) W. C. Cheng & L. K. Fu as a synonym. 73
China: Yunnan, Hengduan Shan, Atuntze, T. T. Y,
7952 (holotype TLF). Distribution
The species epithet refers to the crowded leaves. This species has been included with A. spectabi-
lis (Liu, 1971; Cheng & Fu, 1978; Sahni [in part],
Vernacular names 1990), but it is distinct both morphologically and
geographically. It has somewhat smaller cones with
Sikkim fir; gobria, gobra, salla (Nepal); dhunsing slightly exserting bracts and leaves with slightly
(Bhutan) revolute margins, which are arranged more radially
around the shoots, especially on coning branches. In
Description these characters it is intermediate between A. spec-
tabilis and A. delavayi, the latter a species occurring 79
Trees to 5060 m tall, d.b.h. to 2.5 m; trunk monopo- from NE India to Yunnan, China, and also disjunct
dial, straight, columnar, terete; crown broad, pyra- in N Viet Nam. Rushforth (2009) described a new
midal or columnar, in old trees flat topped. Bark species, A. fordei, from S Xizang [Tibet] ocurring
soon scaly, grey, in old trees fissured towards base, on the northern side of the Himalayan crest in the
breaking into large plates. Branches of first order Yarlung Zangbo drainage. Apart from its flat, not
thick, long, assurgent near the top, horizontally revolute, leaf margins, it is similar to A. densa and
spreading, or lower branches bent down; branches is here treated as synonymous. It thus appears that
of second order assurgent. Branchlets thick in lead- at high elevation in the Himalaya, we find from west
ing shoots, lateral shoots firm but slender, yellow- to east, with partly overlapping distributions, first
ish to reddish brown, becoming grey, prominently A. spectabilis, then A. densa, and finally A. delavayi,
ridged and grooved, glabrous or with slight pubes- which extends into the adjacent mountains of west-
cence in grooves; leaf scars circular ovate. Vegetative ern China and as far as Viet Nam.
buds ovoid-conical, 8 6mm, resinous; bud scales
ovate, obtuse, orange brown and persistent for Distribution
several years. Leaves spirally arranged, more or
less pectinate, on leading and coning shoots more E Himalaya: from E Nepal to the Great Bend of the
radially arranged, with the upper leaves curved Brahmaputra.
forward, (1.5)24(5) cm long, 1.52.5mm wide, TDWG codes: 36 CHT 40 ASS-AS EHM-AP EHM-BH
twisted or curved at base, linear, straight or slightly EHM-DJ EHM-SI NEP
curved, flattened; margins slightly revolute; upper
surface of leaves light or mid green, lower surface Ecology
with two white bands; apex emarginate. Stomata
in two bands separated by a midrib below. Pollen Abies densa occurs in the high mountains of the
cones lateral, radially arranged around the shoot, eastern Himalaya, from 2450 m to 4000 m a.s.l., on
24.5cm long, yellow with purplish blue microspo- rocky, often steep slopes in the cloud belt, where it
rophylls. Seed cones lateral, erect, sessile or on very grows on a variety of alpine lithosols. The climate
short peduncles, (ovoid-)cylindrical with obtuse is extremely wet, with well marked monsoons and
or umbilicate apex, 812cm long, 45.5cm wide, an annual precipitation of more than 2000mm. The
purplish blue when immature, becoming blackish summers are relatively warm, the winters are cold
purple with some brown when ripe. Seed scales fla- at high altitudes and bring heavy snowfall. The spe-
bellate-cuneate, length width at mid-cone 1.52 cies occurs in a wide altitudinal range from mixed
22.5 cm; surface smooth or slightly striated, deciduous coniferous forest at lower elevations to
pubescent on exposed parts; upper margin entire, stands with Betula utilis at tree line. Deciduous trees
incurved; base pedicellate. Bracts ligulate-spathu- are e.g. Acer caudatum, A. pectinatum, Prunus spp.,
late, 22.5cm long, slightly or not exserted. Seeds Sorbus spp. and many large Rhododendron spp. Most
cuneate, 8 4mm, brown; seed wings cuneate, 10 of these disappear above 3000 m to make place for
5mm, brown. conifers. Picea spinulosa and Tsuga dumosa occur
generally in a belt below Abies; Larix griffithiana and/ 45mm long, very resinous; bud scales dark brown
or Juniperus squamata above it, the latter at tree line. (but covered with yellowish resin), persisting 23
years. Leaves spirally arranged, more or less pecti-
Conservation nate, curved slightly backward or forward, on con-
ing shoots the same or weakly assurgent, (1.4)
IUCN: LC 23.5(4.5) cm long, 11.6mm wide, twisted at
base, narrowly linear, tapering towards an acutish
Uses or obtuse apex, (dark) green or light green; none or
with a few lines of stomata above, two bands sepa-
Sikkim fir is a timber tree of importance in the rated by a midrib below. Pollen cones lateral, in leaf
80 eastern Himalaya, where it is used in construction axils, 12 (?) cm long (according to Liu, 1971, 5mm,
(house building), in particular for interior work such but likely longer when mature), with resinous peru-
as floor boards, ceilings and stairs, while shingles are lar scales; microsporophylls red. Seed cones lateral,
used for roofing. This species has been introduced erect, on very short peduncles, cylindrical with
in Europe only relatively recently from Bhutan, obtuse apex, 5.510cm long, 34.5cm wide; pale
Nepal and Sikkim. It is still uncommon in gardens brown or ochraceous when immature, ripening to
and parks but has more attractive characters than A. pale brown; cone rachis persistent, narrowly conical.
spectabilis, with very white leaf undersides and deep Seed scales flabellate-cuneate, usually wider than
purplish blue, nearly black seed cones. It undoubt- long, length width at mid-cone 1.52 22.8cm;
edly requires ample precipitation, judging from its surface smooth, pubescent on exposed parts; upper
natural habitat. margin entire, slightly reflexed; base pedicellate.
Bracts spathulate, with short cusps, 11.5cm long,
included. Seeds cuneate-oblong, 78mm long, yel-
Abies durangensis Martnez, Anales Inst. Biol. Univ. lowish brown; seed wings cuneate to more or less
Nac. Mxico 13 (2): 621. 1942. rounded, 810 68mm, pale yellowish.
Etymology Distribution
The species epithet refers to the Mexican state of Mexico: Chihuahua, Coahuila, Durango, N Jalisco,
Durango. Sinaloa.
TDWG codes: 79 MXE-CO MXE-CU MXE-DU
Vernacular names MXN-SI MXS-JA
IUCN: VU (D2)
Abies neodurangensis Debreczy, Rcz & Salazar,
Phytologia 78 (3): 7. 1995.
Abies fabri (Mast.) Craib, Notes Roy. Bot. Gard.
Description Edinburgh 11: 278. 1919.
Farges fir; Bashan lengshan (Chinese) Abies fargesii occurs in the high montane to subal-
pine zones of N Central China, at elevations between
Description 2000 m and 4000 m a.s.l. (A. fargesii var. faxoniana
at elevations between 2600 m and 4000 m, with an
Trees to 40 m tall, d.b.h. to 1.52 m; trunk mono- optimum between 3400 m and 3800 m according to
podial, straight, columnar, terete; crown narrowly Wang, 1961). Soils are mostly grey brown mountain
pyramidal or conical. Bark of young trees smooth, podzols. The climate is cold and moist. At its lowest
finely flaking, grey, of old trees fissured, brown elevation broad-leaved trees (e.g. Fagus engleriana,
grey. Branches of first order massive, short, sparse; Davidia involucrata) are important, but A. fargesii
branches of second order spreading, assurgent, or mostly forms either pure forests or mixed conifer-
lower branches pendant. Branchlets slender, firm, ous forests with among other species Picea purpurea,
reddish brown, purplish or mahogany (variable), P. asperata, P. neoveitchii, P. brachytyla, Larix potani-
shallowly grooved, glabrous, minutely pubescent (in nii, Abies chensiensis, A. recurvata, Tsuga chinensis
grooves) or densely pubescent on 1st year shoots; leaf and Taxus chinensis. Some broad-leaved trees are
scars circular. Vegetative buds ovoid-oblong, (4)58 usually present: Betula spp., Populus spp., and many
45mm, slightly resinous; bud scales triangular shrubs: Cotoneaster, Ribes, Spiraea, Rhododendron
ovate, yellowish brown, persistent for several years. and Berberis are among the common genera (except
Leaves spirally arranged, crowded in several overlap- in dense Picea-Abies forest).
ping ranks of unequal length, the lower leaves pecti-
nate, on coning shoots assurgent, (1)1.53(4.5) cm Uses
long, 23.5mm wide, twisted at base, ligulate-linear
or linear, flat or margins slightly revolute; shining Being the most widespread of firs in the high moun-
green above, whitish or glaucous green below; apex tains of western China, this species has been subject
entire, emarginate or bifid, obtuse or acute on cone to extensive exploitation for its timber. The wood,
bearing shoots. Stomata in two bands divided by if of high grade, is used in construction (mainly
a midrib on the lower side of leaves. Pollen cones indoor flooring, framing and joinery), otherwise it
crowded, axillary near shoot apex, small, yellow is applied in the paper pulp industry. Most botanical
with red microsporophylls. Seed cones lateral, erect; collectors active in western China in the first decades
of the 20th century encountered it or its varieties and with flat margins and emarginate apex. Seeds light
the species has been introduced to Europe and the brown, wings pale, tinged purple.
USA from these collections. Most trees still in culti-
vation date from these introductions and are gener- Distribution
ally confined to botanical gardens and arboreta.
China: NW Sichuan, Gansu.
3 varieties are recognized: TDWG codes: 36 CHC-SC CHN-GS
Description Etymology
New shoots densely pubescent with ferruginous The species epithet (Latin: firmus = firm, stable)
short hairs; buds 45cm long, 4mm wide; leaves refers to the rigidity of the leaves.
Vernacular names Distribution
Momi fir; momi, momi-noki (Japanese) Japan: Honshu, Kyushu, Shikoku, Yakushima.
TDWG codes: 38 JAP-HN JAP-KY JAP-SH
Description
Ecology
Trees to 50 m tall, d.b.h. to 2 m; trunk monopo-
dial, massive, straight, columnar, terete; crown Abies firma occurs on hills and in mountains of
open, broad pyramidal, domed or flat topped in old southern and central Japan, at elevations between
trees. Bark of young trees smooth, with horizontal 50 m and 1200 m a.s.l. (commonly 300 m and 1000
86 resin blisters, pinkish grey, in old trees thick, corky, m). The soils are various mountain soils of volcanic
rough, fissured at base. Branches of first order long, origin or alluvial, and mesic. The climate is moist or
spreading horizontally or, especially in young trees, wet, cool in the north of its range and warm tem-
ascending; branches of second order thick, spread- perate in the south, with annual precipitation above
ing horizontally or ascending. Branchlets firm, yel- 1000mm. This species is a constituent of mixed
lowish grey, buff grey brown or light brown, grooved, forests (rarely in pure stands on dry sites) with
usually quite glabrous or with fine pubescence in e.g. Fagus crenata, F. japonica, Castanea crenata,
grooves of 1st year shoots; leaf scars circular or angu- Carpinus laxifolia, Quercus spp., Tsuga sieboldii,
lar. Vegetative buds ovoid to conical, maximal 10 Pinus parviflora, P. densiflora, Pseudotsuga japonica,
5mm, not or only slightly resinous; bud scales broad Abies homolepis, Cryptomeria japonica, Sciadopitys
conical, pale (reddish) brown with green margins, verticillata, Chamaecyparis obtusa, Torreya nucifera
persisting several years. Leaves spirally arranged, and Picea jezoensis subsp. hondoensis.
pectinate in 23(4) ranks of different length, on
coning shoots spreading, the upper leaves assurgent, Conservation
(1)1.53.5(5) cm long, 24mm wide, narrowed
and curved or twisted at base, linear, flattened; lus- IUCN: LC
trous (light) green above, whitish green below; apex
obtuse or emarginate (bifid in young trees). Stomata Uses
in two broad bands separated by a midrib below,
none or a few in a central groove above. Pollen cones Momi is the most common and widespread fir in
solitary in leaf axils, pendulous, 2.53cm long, yel- southern Japan and there regarded as an impor-
low. Seed cones lateral, erect; peduncles short, with tant timber tree. Its wood is light, soft and straight-
many scales at base; shape ovoid-oblong or coni- grained and easily worked, but requires careful
cal, with obtuse apex, 815cm long, 35cm wide; seasoning to prevent warping. It is used for carpen-
yellowish green or green with yellow bracts when try making indoor framing, flooring, joinery, crates,
immature, maturing to greyish green and becoming boxes etc., but the greatest quantities of its timber are
yellowish brown when ripe; cone rachis persistent, converted to paper pulp. In plantation forestry it is
narrowly conical, brown. Seed scales broad flabellate, only common in Japan, where old growth stands of
length width at mid-cone 22.5 2.83.2cm; sur- this large fir have mostly been logged. Elsewhere, it
face smooth or longitudinally striated, glabrous or is only used as an ornamental tree or planted in col-
sparingly pubescent on exposed parts; upper margin lections in arboreta and botanic gardens, requiring
thinner than the rest of the scale, incurved, entire or a climate with mild winters and abundant rainfall.
obscurely denticulate; base short pedicellate. Bracts
oblanceolate, 2cm long, exserted, especially on
lower part of cone, straight. Seeds obovate-cuneate,
68mm long, light brown; seed wings broad cune-
ate, 1015mm long, yellowish brown.
Abies forrestii Coltm.-Rog., Gard. Chron., ser. 3, 65: the cusps exserted, with straight or recurved cusps.
150. 1919. Seeds obovate, ca. 8mm long, lustrous brown; seed
wings cuneate-dolabriform, 10 8mm, light brown.
Etymology
Taxonomic notes
This species was named after the English plant
hunter George Forrest (18731932). Abies chengii Rushforth was described from cul-
tivated trees in the British Isles, derived from seed
Vernacular names collected by George Forrest somewhere in Yunnan
and was accepted as a species in my book Pinaceae
Forrest fir; chuandian lengshan (Chinese) (Farjon, 1990). It is more probably another variety 87
(or a mere form not deserving a botanical name) of
Description A. forrestii. A hybrid origin with this species and A.
chensiensis subsp. salouenensis has also been sug-
Trees to 40 m tall, d.b.h. to 11.5 m; trunk monopo- gested, but without supporting evidence. Abies for-
dial, straight, columnar, terete; crown broad conical, restii is evidently a variable species.
flat topped in old trees. Bark in young trees smooth,
brown grey, in old trees dark brown, fissured at base. Distribution
Branches of first order long, spreading horizontally,
ascending near the top; branches of second order China: SW Sichuan, NW Yunnan, SE Xizang [Tibet].
assurgent. Branchlets thick, purplish, reddish or TDWG codes: 36 CHC-SC CHC-YN CHT
orange brown, often turning grey, smooth or finely
grooved, glabrous or (ferruginous) pubescent; leaf Ecology
scars circular or oval. Vegetative buds globular to
ovoid, 410 37mm, very resinous; bud scales This species (and its varieties) occurs in the high
ovate, red brown (covered with white resin), persist- mountains of SW China at elevations between 2400
ing several years. Leaves spirally arranged, dense, m and 4300 m a.s.l. (commonly 30004000 m), on
covering shoot in several ranks, parting in the grey-brown mountain podzols. The climate is cold
middle, assurgent on coning shoots, (1.5)23(4) and wet, annual precipitation ranges from 1000mm
cm long, 22.5mm wide, curved or twisted at base, to 2000mm. The species forms forests in pure
linear or ligulate-linear, flattened; margins flat or stands near the tree limit, or is mixed with Picea liki-
slightly revolute; lustrous dark green or bluish green angensis, Larix potaninii, Tsuga dumosa and some
above, two white (sometimes tending to greenish broad-leaved trees, e.g. Betula albo-sinensis, Acer
white) bands below; apex emarginate, or acumi- spp. and Sorbus spp. at lower elevations. An erica-
nate on coning shoots. Stomata in two bands sepa- ceous lower shrub layer with Rhododendron spp. is
rated by a green midrib below. Pollen cones lateral, often prominent.
34.5cm long, yellowish, with purple microsporo-
phylls. Seed cones lateral, erect on short peduncles, Uses
thick, barrel shaped or cylindrical, with obtuse or
retuse apex, 610(14) cm long, 45(6) cm wide, Forrest fir and and its several varieties occur at high
purplish blue with blue bracts when immature, rip- altitudes, often up to the tree line and consequently
ening to purplish brown or dark brown; cone rachis only yield timber suitable for saw mill process-
persistent, massive, cylindro-conical or fusiform, ing from larger trees at its lowest altitudinal range.
purplish brown. Seed scales cuneate-obovate, length Exploitation has (at least officially) ceased with
width at mid-cone 2 1.8cm; surface smooth or Chinese forest conservation law now prohibiting
somewhat rough, puberulent; upper margin entire logging in old growth forest in the western prov-
or slightly erose; base pedicellate. Bracts oblong or inces. Having been collected on numerous occasions
broadly cuneate-spathulate, 23cm long, with long by the famous European plant hunters of the early
cusps (10mm), much exserted, included or only 20th century it was introduced to Europe and the
United States where it is still quite common in arbo- Abies ferreana Bordres & Gaussen var. longibrac-
reta and private large gardens. Most trees labeled A. teata L. K. Fu & Nan Li, Novon 7 (3): 261. 1997.
delavayi actually belong to this species (A. delavayi
has narrow leaves with revolute margins and dark Description
violet-blue or purplish black seed cones). Renewed
collecting in recent decades has brought in some Shoots pubescent. Resin canals in leaves medial.
new stock on a limited scale. Abies forrestii is one of Seed cones 68cm long.
the most attractive species of fir in horticulture and
deserves to be made much more widely available. Distribution
Perhaps China could do some alternative business
88 now that they have ceased logging? China: NW Yunnan, SE Xizang [Tibet].
TDWG codes: 36 CHC-YN CHT
4 varieties are recognized:
Conservation
Abies forrestii Coltm.-Rog. var. forrestii. Abies dela-
vayi Franch. var. forrestii (Coltm.-Rog.) A. B. Jacks., Logging, especially when followed by grazing (as at
in Chittenden, Cult. Conif.: 245. 1932. Type: China: Zhongdian where there are yaks), by fire and by the
Yunnan, Lijiang Shan, eastern slopes at 27 25 N, G. planting of other conifer species present an identi-
Forrest 6744 (holotype E). fiable threat. A past reduction of ca. 50% seems to be
a reasonable estimate.
Abies chengii Rushforth, Notes Roy. Bot. Gard. IUCN: EN (A2cde)
Edinburgh 41: 333. 1983; Abies forrestii Coltm.-Rog.
var. chengii (Rushforth) Silba, Phytologia 68: 17. 1990.
Abies forrestii Coltm.-Rog. var. georgei (Orr)
Description Farjon, Pinaceae (Regnum Veg. 121): 59. 1990. Abies
georgei Orr, Notes Roy. Bot. Gard. Edinburgh 18: 1,
Shoots glabrous. Resin canals in leaves marginal, t. 236. 1933; Abies delavayi Franch. var. georgei (Orr)
small. Bracts slightly exserted, or only the cusps Melville, Bull. Misc. Inf. R.B.G. Kew 1958: 533. 1959.
exserted, or sometimes entirely included. Type: China: Yunnan, Jinsha-Mekong Divide, G.
Forrest 22547 (holotype E).
Distribution
Description
China: SW Sichuan, NW Yunnan, SE Xizang [Tibet].
TDWG codes: 36 CHC-SC CHC-YN CHT Shoots densely pubescent with ferruginous, short
hairs; leaves 1.53cm long; margins slightly revolute.
Conservation Bracts of seed cones exserted, broad and gradually
tapering to a short cusp, with erose-denticulate, light
IUCN: LC brown margins.
Abies chayuensis W. C. Cheng & L. K. Fu, Acta China: SW Sichuan, NW Yunnan, SE Xizang [Tibet].
Phytotax. Sin. 13 (4): 83. 1975; Abies forrestii Coltm.- TDWG codes: 36 CHC-SC CHC-YN CHT
Rog. var. chayuensis (W. C. Cheng & L. K. Fu) Silba,
Phytologia 68: 16. 1990.
Conservation often open and irregular on exposed sites, broader
in solitary trees. Bark of young trees smooth, brown,
IUCN: LC with numerous horizontal resin blisters, becoming
rough, scaly and grey in old trees. Branches of first
order long, spreading horizontally or ascending near
Abies forrestii Coltm.-Rog. var. smithii Vigui & the top; branches of second order slender, spreading
Gaussen, Trav. Lab. Forest. Toulouse T. 1 (2, 1): 177. horizontally or assurgent. Branchlets slender, stiff,
1929. Abies delavayi Franch. var. smithii (Vigui & pale yellowish brown, becoming darker brown in the
Gaussen) T. S. Liu, Monogr. Gen. Abies: 143. 1971; second year, slightly grooved, densely and rather per-
Abies georgei Orr var. smithii (Vigui & Gaussen) sistently pubescent with short, reddish hairs; leaf scars
W. C. Cheng & L. K. Fu, Acta Phytotax. Sin. 13 (4): circular. Vegetative buds broadly ovoid, 4 3mm, 89
63. 1975. Type: China: Yunnan, Lijiang Shan, E very resinous; bud scales dark reddish brown, but
slope, J.F. Rock 10673 (holotype A). covered with yellowish resin, persistent several years.
Leaves spirally arranged, the lower ones pectinate, the
Description upper ones spreading upward and forward, incurved
and assurgent especially on coning shoots, 12(2.3)
Shoots pubescent. Bracts of seed cones exserting, cm long, 22.2mm wide, twisted or curved at base,
broad and abruptly narrowing to a long cusp. linear or falcate linear, widest near the obtuse or acute
(rarely emarginate) apex, dark green above, two whit-
Distribution ish bands below. Stomata in two bands separated by a
midrib below, a few in the central groove above. Pollen
China: NW Yunnan. cones crowded on the lower side of branches, pendu-
TDWG codes: 36 CHC-YN lous, 1cm long, yellow, with red microsporophylls.
Seed cones lateral, erect, often crowded, on short
Conservation peduncles, oblong-conical, with narrowed, obtuse
apex, 48cm long, 2.54cm wide, dark purple with
IUCN: NT yellowish bracts when immature, ripening to purplish
brown with pale brown bracts; cone rachis persistent,
narrowly conical, dark brown. Seed scales broadly fla-
Abies fraseri (Pursh) Poir. in Lamarck, Encycl. bellate to reniform, length width at mid-cone 0.8
Suppl. 5: 35. 1817. Pinus fraseri Pursh, Fl. Amer. Sept. 1.2 11.5cm; surface smooth, puberulent on exposed
2: 639. 1814; Abies balsamea (L.) Mill. subsp. fraseri parts; upper margin entire and incurved; base long
(Pursh) E. Murray, Kalmia 12: 18. 1982. Type: not pedicellate. Bracts oblong or obovate-oblong, with
designated. obcordate apices and short cusps, 1.52cm long,
much exserted and reflexed, covering most of the
Etymology seed scales. Seeds obovate-cuneate, 56mm long,
dark purplish black; seed wings obliquely cuneate or
This species was named after John Fraser (17501811) dolabriform, 56 5mm, purplish brown.
who collected plants in eastern North America,
among which was this fir. Distribution
Description Ecology
Trees to 15 m tall, d.b.h. to 0.5 m; trunk monopodial, On the highest slopes and summits of the
straight, columnar, terete; crown narrowly conical, Appalachian Mountains, between 1200 m and 2150 m
a.s.l., usually best developed on N-facing slopes. Abies grandis (Douglas ex D. Don) Lindl., Penny
The soils are commonly podzolized and moderately Cyclop. 1: 30. 1833. Pinus grandis Douglas ex D.
acid. The climate is humid, with cool summers and Don, in Lambert, Descr. Pinus, ed. 8, 2: p. s.n. inter
cold winters with heavy snowfall, annual precipita- 144 et 145 Type: not designated.
tion varies between 850mm and 2000mm. Fraser
fir occurs in scattered populations, sometimes pure Abies excelsior Franco, Bol. Soc. Brot., ser. 2, 23: 162.
at the highest elevations, but more often mixed with 1949.
Picea rubens and Betula papyrifera above 1500 m, at Abies grandis (Douglas ex D. Don) Lindl. var. ida-
lower elevations also with Tsuga caroliniana, Betula hoensis Silba, Phytologia 68: 19. 1990.
alleghaniensis, Sorbus americana, Acer saccharum
90 and Fraxinus caroliniana. Ericaceae and various Etymology
herbs are common in the understorey, often thick
moss carpets (Hylocomium splendens) cover the for- The species epithet is a reference to the great stature
est floor. this species can attain in its natural habitat.
The disjunct populations of this fir, restricted to the Grand fir, Lowland fir, Giant fir
mountain tops and their north-facing slopes of the
southern Appalachians, are susceptible to destruc- Description
tion by windfall and fire. However, by far the most
damaging agent is an insect, the Balsam woolly adel- Trees to 80100 m tall, d.b.h. to 23 m; trunk mono-
gid (Adelges piceae), discovered in 1957 in Abies fra- podial, straight, columnar, terete; bole often free
seri on Mt. Mitchell. This alien pest has spread quickly of branches to considerable height; crown nar-
to all populations causing massive dieback through rowly conical. Bark of young trees thin, smooth
impairment of translocation flow in the cambium. and grey brown, with many resin blisters, braking
Millions of trees had died by the 1980s and only up into many small plates in old trees. Branches of
one substantial population (Mt. Rogers, Virginia) first order long, spreading horizontally, the lower
remained largely unaffected. Methods to control this ones curved downward; branches of second order
introduced insect are still being researched but none spreading, or ascending near the end of the main
have been fully effective; some small scale protection branches. Branchlets slender, firm, olive green to
can be provided by chemical insecticides. reddish brown, with narrow, straight ridges between
IUCN: EN [B2 (ii, iv, v)] the leaves, minutely pubescent, but glabrous by the
3rd year; leaf scars small, circular. Vegetative buds
Uses globose, 1.52 1.5mm, slightly resinous; bud scales
triangular, obtuse, red-brown, persisting 23 years.
The remaining stands of Fraser fir have very limited Leaves spirally arranged, pectinately spreading in
commercial value as timber trees. The most impor- a more or less horizontal plane, the upper ranks
tant use is growing this species for Christmas trees; shorter than the lower, on coning shoots assurgent,
it is considered the best conifer available in the USA (2)35(6) cm long, 23mm wide, strongly twisted
for this purpose. It has a natural Christmas tree at base, narrowly linear, flattened with slightly revo-
shape and retains its fragrant, dark green leaves lute margins, dark glossy green above, greenish white
well for this indoor use. It is also widely used as an below; apex emarginate or obtuse on coning shoots.
ornamental tree for gardens with several cultivars Stomata only on the abaxial (lower) surface in two
named. At least in the UK it does not usually have a bands separated by a midrib. Pollen cones lateral on
very long life as a garden tree. the lower sides of branches, axillary, 1.21.8cm long,
yellowish green. Seed cones lateral, erect; peduncles
very short (cones nearly sessile); shape oblong-cylin-
drical, with obtuse apex, (5)712cm long, 34cm
wide, light green or purple tinged when immature, and an excellent source of pulpwood. For construc-
ripening to dull grey-brown, usually very resinous; tion timber it is considered less desirable due to its
cone rachis persistent, narrowly conical, brown. relative weakness and limited durability. In the Pacific
Seed scales broad flabellate, length width at mid- Northwest young trees are valued as Christmas trees
cone 22.5 2.53cm; surface smooth, in immature because they tend to grow up very symmetrically
cones pubescent; upper margin entire, incurved; and have lustrous green foliage. Grand fir is com-
base pedicellate. Bracts short, rectangular, with very monly grown as an amenity tree in large gardens and
small cusps, 11.5cm long, entirely included. Seeds city parks and, as another David Doulas introduc-
cuneate, 8 5mm, pale brown; seed wings cuneate- tion, it was planted in nearly all landscape gardens
dolabriform, 1015mm long, pale brown with a pur- laid out in the 19th century in Europe, where some
ple tinge. trees have now attained impressive sizes. In horti- 91
culture it is much in use and a substantial number
Distribution of cultivars have been selected for garden planting.
Abies hickelii Flous & Gaussen, Trav. Lab. Forest. Taxonomic notes
Toulouse T. 1 (1, 17): 1. 1932.
In a recent paper Strandby et al. (2009) presented a
Etymology morphometric study of the genus Abies in Mexico
and Central America. They proposed to reduce A.
This species was named after the French botanist hickelii to a subspecies of A. religiosa on the basis
and dendrologist R. Hickel. of their results. This paper came too late for these
results to be thoroughly considered (or tested) and
Vernacular names therefore this new taxonomy is merely noted here.
The type collection, Debreczy et al. DAPC 40323 Abies holophylla Maxim. var. aspericorticea Y. Y. Sun,
(holotype BP, isotypes A, E, K, MEXU) was gathered Bull. Bot. Res. (China) 25 (3): 264. 2005.
4.5 km N of the village of Metepec. The number of
resin ducts in the leaves, given as 57 in the proto- Etymology
logue, is highly unusual in the genus, but A. hickelii
has 412. Abies hidalgensis could be nothing more The species epithet means entire leaf and refers to
than a variety of A. hickelii with included bracts in the the undivided leaf apex.
seed cones. It is only known from the type specimens
deposited in the herbaria cited above. More material Vernacular names
is needed to establish its taxonomic position, the spe-
cies is here given the benefit of the doubt. Needle fir, Manchurian fir; shansong (Chinese)
Distribution Description
Mexico: Hidalgo (canyon near village of Metepec). Trees to 3050 m tall, d.b.h. to 11.5 m; trunk mono-
TDWG codes: 79 MXE-HI podial, straight, columnar, terete; crown shape broad
pyramidal, young trees with narrowing top. Bark
Ecology of young trees smooth, grey to buff orange, in old
trees shallowly fissured, scaly and brown. Branches
This species occurs in a steep canyon in the Sierra of first order long, spreading horizontally; branches
Madre Oriental at an altitude of ca. 20002300 m of second order spreading horizontally or assurgent.
Branchlets stout, firm, yellowish to grey-brown or snow. At higher elevations or in the NE of its range
buff orange, with ridges and grooves between the it forms pure stands, or more commonly mixed
leaves, glabrous, or with only minute pubescence coniferous forests with Pinus koraiensis, especially
in grooves of 1st year shoots; leaf scars oval circular, in the coastal mountains near the Sea of Japan. In
light coloured. Vegetative buds ovoid-conical, 48 other areas it is a constituent of the northern mixed
35mm, more or less resinous; bud scales triangular coniferous deciduous forests, with Abies nephrol-
with blunt apices, prominently keeled, light brown epis, Picea obovata, Larix gmelinii (var. olgensis) and
or reddish brown, persisting several years. Leaves broad-leaved trees, such as Populus spp., Quercus
spirally arranged, radial and assurgent on lead- mongolica, Fraxinus mandshurica, Ulmus spp., and
ing and coning shoots, subdistichous and/or more Betula ermanii.
96 or less pectinate in shaded shoots, 24.5cm long,
22.5mm wide, twisted or curved at base, linear, Conservation
tapering towards an acute or mucronate apex, flat-
tened, lustrous light green above, pale whitish green IUCN: NT
with a green midrib and margins below. Stomata
in two bands divided by a midrib below, none or a Uses
few near apex above. Pollen cones lateral, clustered,
11.5cm long, yellow with reddish microsporo- Manchurian fir is a valuable timber tree in its native
phylls. Seed cones lateral, erect; peduncles 0.51cm, area and is exploited mainly in managed forests in
scaly; shape oblong-cylindrical, with obtuse or trun- much of the region. Most of its wood today is used
cate apex, 614cm long, 34.5cm wide, pale green for plywood and veneer as it is evenly grained, light
when immature, sometimes with a purple tinge, and easily worked. Minor uses are for soundboards
maturing to yellowish green and ripening to light in musical instruments, boxes and sometimes for
yellowish brown; cone rachis persistent, narrowly joinery. In Europe it was introduced around 1905,
conical, brown. Seed scales flabellate, length width but, although perfectly hardy and ornamental,
at mid-cone 2 3cm; surface smooth, puberulent remains an uncommon tree mainly seen in arboreta
on exposed parts, often very resinous; upper mar- and botanic gardens of the northern hemispheres
gin entire, slightly incurved; base pedicellate. Bracts cooler regions.
oblong-spathulate, with small cusps, 0.81cm long,
included. Seeds cuneate, ca. 6 4mm, light brown;
seed wings cuneate, with a truncate, oblique end, ca. Abies homolepis Siebold & Zucc., Fl. Japon. 2 (2): 17,
13 10mm, light brown. t. 108. 1842. Fig. 8
Distribution Etymology
NE Asia: from mountains N of Vladivostok to South The species epithet (Greek/Latin: homo = equal, the
Korea, also in Heilongjiang, Jilin and Liaoning Prov., same; lepis = scale) refers to the similar length of
China. seed scales and bracts.
TDWG codes: 31 PRM 36 CHM-HJ CHM-JL
CHM-LN 38 KOR-NK KOR-SK Vernacular names
Ecology IUCN: NT
Etymology Distribution
Taiwan fir; Taiwan lengshan (Chinese) A high mountain species, occurring between 2400
m and 3800 m a.s.l. in the central high mountains
Description of Taiwan, on grey brown podzolized soils and also
on mountain yellow earth, both acid and usually
Trees to 1620 m tall, d.b.h. to 0.51 m; trunk mono- rocky. The climate is temperate, super humid: above
podial, straight, terete, in some old trees bent, but humid subtropical foothills the annual precipitation
otherwise erect; crown broad pyramidal, on moun- exceeds 4000mm, with maxima up to 10,000mm,
tain tops irregular and open. Bark of young trees making the Taiwanese central high mountains one
smooth, soon flaking, light grey or yellowish grey, in of the wettest mountain ranges in the world. There
old trees becoming rough and scaly, greyish brown. are some pure forests on N and NE slopes at these
Branches of first order heavy and long, spreading high elevations (3200 m to 3600 m a.s.l.), or the
horizontally or curved; branches of second order species occurs mixed with scattered Pinus arman-
spreading or assurgent. Branchlets stout, firm, dii var. mastersiana, Tsuga chinensis var. chinensis,
pale yellowish brown, with prominent ridges and Picea morrisonicola, and with Juniperus squamata
grooves, light brown pubescence in grooves, on cone var. morrisonicola at the upper limit of Abies. At
bearing shoots pubescence red brown; leaf scars lower elevations the forest becomes progressively
circular. Vegetative buds conical-ovoid, 45mm more mixed with broad-leaved trees, e.g. Acer insu-
long, very resinous; bud scales triangular, keeled, lare, Trochodendron aralioides, Quercus semecarpi-
purple-brown, persisting several years. Leaves spi- folia subsp. glabra, Ilex bioritsensis, and Eurya spp.
rally arranged, spreading radially, pectinate below Other conifers in this belt are Tsuga chinensis var.
shoot, shorter and assurgent above shoot, cover- chinensis, which becomes more abundant than
ing it entirely, on coning shoots assurgent, (0.8)1 Abies kawakamii between 2400 m and 3000 m a.s.l.,
2.2(2.5) cm long, 1.52mm wide, slightly twisted Pseudotsuga sinensis, and Chamaecyparis obtusa var.
or curved at base, linear-falcate, flattened, lustrous formosana, which is more abundant below 2400 m
green above, whitish green below; apex emarginate (Liu, 1971).
Conservation the top. Branchlets slender, firm, yellowish grey or
grey-green, shallowly grooved between the leaves,
Logging of this species, which occurred mainly dur- sparsely pubescent in the grooves, soon glabrous;
ing the period of Japanese occupation (18951945), leaf scars circular. Vegetative buds subglobose, 45
has ceased almost completely and substantial popu- 34mm, very resinous; bud scales obtuse, membra-
lations now occur within national parks and other nous, brown, persisting several years. Leaves spirally
reserves. Its limited distribution and occurrence in arranged, radially spreading, the upper leaves shorter
a mosaic with subalpine bamboo grassland makes and curved upward or recurved, especially so on
it vulnerable to fires that could be caused by much coning shoots, (0.8)12(2.2) cm long, 22.5mm
increased tourism. wide, slightly twisted or curved at base, linear-
IUCN: NT spathulate (often widest near the apex), flattened, 99
with slightly revolute margins, lustrous dark green
Uses above, two white bands and green midrib and mar-
gins below; apex emarginate, obtuse or sometimes
The timber of this species was formerly exported acute. Stomata none or a few near the apex above, in
to Japan, where it was used for general carpentry. It two broad bands separated by a midrib below. Pollen
is little used for this purpose today in Taiwan. This cones lateral, clustered around shoot, 1cm long, yel-
species was introduced to England in 1930 and is lowish with scarlet microsporophylls. Seed cones lat-
occasionally found in arboreta in Europe and North eral, erect, often crowded together, numerous, even
America, but remains uncommon in cultivation. on the lower branches; peduncles 0.40.6cm; shape
cylindrical, with obtuse, umbilicate or papilliform
apex, 47cm long, 2.53cm wide, purple or violet
Abies koreana E. H. Wilson, J. Arnold Arbor. 1: blue with greenish bracts when immature, maturing
188. 1920. Type: South Korea: Cheju-do, Halla-san, to purplish brown, with light brown bracts, ripening
[Hallai-san Quelpart Island], E. H. Wilson 9486 to dull brown or purplish brown; cone rachis per-
(holotype A). Fig. 10 sistent, narrowly conical, dark brown. Seed scales
reniform, or wing-shaped (Liu, 1971, p. 45), length
Abies koreana E. H. Wilson f. nigrocarpa Hatus., width at mid-cone 11.2 1.21.6cm; surface smooth,
Bull. Kyushu Univ. Forest. 5: 40. 1934. pubescent; upper margin erose, incurved; base pedi-
cellate. Bracts broadly spathulate-obcordate, 1.2
Etymology 1.5cm long, exserted, (strongly) reflexed. Seeds cune-
ate, 46mm long, light or dark brown, with a purple
The species name denotes its origin, Korea where tinge; seed scales cuneate-dolabriform, 46mm
Ernest Wilson discovered it. long, light brown, tinged with purple.
Description Ecology
Trees to 6070 m tall, d.b.h. to 23 m; trunk mono- Abies magnifica occurs in the Canadian Life Zone of
podial, massive, straight, columnar, terete; crown high mountains, between 1400 m and 2700 m a.s.l.
narrowly conical, in old trees open. Bark of young (to 3000 m in the south of its range); commonly on
trees smooth, with resin blisters, grey or light pur- soils of granitic (Sierra Nevada) or basaltic (Cascade
plish grey, in old trees thick, rough, deeply fissured Range) origin, which have been altered by glaciation
below, brown. Branches of first order relatively and are usually slightly acid. The climate is character-
short, spreading, the lower bent downward or pen- ized by short, warm and dry summers and long, cold
dant; branches of second order spreading horizon- winters with much snow. Annual precipitation var-
tally, pendant at last. Branchlets slender, strong but ies between 750mm and 1500mm (80 % as snow).
pliable, light brown or greenish, with ridges and This species forms pure stands in some places, but
grooves between leaves, for about two years densely more often it is a constituent of the mixed conifer-
pubescent with reddish brown hairs; leaf scars circu- ous forest type with e.g. Pinus spp., Abies concolor,
lar. Vegetative buds ovoid globose or more conical, A. procera, Pseudotsuga menziesii, Calocedrus decur-
small, hidden by terminal short leaves, not resinous; rens, Juniperus occidentalis, and at higher eleva-
bud scales triangular, keeled, brown puberulent, per- tions Abies lasiocarpa and Tsuga mertensiana subsp.
sisting 23 years. Leaves spirally arranged, spreading grandicona. Common shrubs are e.g. Ceanothus cor-
laterally, but curved inward above shoot, on coning dulatus, Chrysolepis sempervirens and Arctostaphylos
shoots assurgent and strongly curved, concealing nevadensis.
shoot, 23.5cm long, 1.31.6mm wide, (abruptly)
curved at base, narrowly linear, curved, only slightly Uses
flattened, no groove above, glaucous grey green on
both sides; apex obtuse or acute. Stomata in two sep- California red fir grows to large dimensions with
arate rows of lines above, in two bands separated by a extremely straight boles and has a high wood pro-
midrib below. Pollen cones lateral, crowded, pendu- duction per ha even in natural, unmanaged stands.
lous, 1.52cm long, with scarlet microsporophylls. It is therefore increasingly valuable as a timber tree
Seed cones lateral, erect, short pedunculate or ses- used for general construction and plywood. This spe-
sile, broad conical or barrel shaped, with truncate or cies is also valued as a Christmas tree, both grown in
umbilicate apex, (10)1320cm long, (5)710cm natural stands and in plantations. It is relatively rare
wide, purplish green when immature, maturing to in amenity plantings with few cultivars known; most
yellowish brown or greenish brown, ripening to light existing planted trees date from the heydays of land-
brown; cone rachis persistent, narrowly conical, scape conifer plantings in the 19th century.
dark brown. Seed scales cuneate-flabellate, length
width at mid-cone (2.5)34 (2.5)34cm; 2 varieties are recognized:
surface smooth, densely pubescent with yellowish
Abies magnifica A. Murray bis var. magnifica. Type: zone, that shows exserted bracts. Artificial crossing
USA: California, Sierra Nevada, W. Lobb 441 (holo- experiments indicate the possiblity of introgression
type K). via pollen from A. procera into A. magnifica.
Description Distribution
Seed cones 1420cm long, 710cm wide; bracts USA: from Lassen Peak in California to Crater Lake
included. in Oregon.
TDWG codes: 73 ORE 76 CAL
Distribution
Conservation 103
USA: California, W Nevada, SW Oregon.
TDWG codes: 73 ORE 76 CAL NEV IUCN: LC
Conservation
Abies mariesii Mast., Gard. Chron., ser. 2, 12: 788.
IUCN: LC 1879. Type: Japan: Honshu, Awomori Pref., Nikko-
san, C. Maries 73 (holotype K). Pl. 2
plate 2. Abies mariesii. 1. Habit of tree. 2. Foliage branch. 3. Seed cone. 4, 5. Seed scales with bract (4),
with seeds (5). 6. Rachis of seed cone. 7. Leaves.
and curved, flattened, grooved above, with slightly Conservation
revolute margins and emarginate apex (obtuse on
coning shoots), lustrous dark green above, 2 white IUCN: LC
bands below separated by green midrib. Stomata
in two bands separated by a midrib on lower sur- Uses
face only. Pollen cones lateral, axillary, pendulous,
1.52cm long, yellowish. Seed cones lateral, short This species of fir has little value as a timber tree
pedunculate or sessile, ovoid-oblong, with obtuse because it grows at high altitude and mostly in inac-
apex, 49cm long, 24.5cm long, violet-blue, rip- cessible localities. In horticulture it is rather uncom-
ening to dark blackish purple (brown inside); cone mon despite its attractive dark green foliage leaves
rachis persistent, narrowly conical, blackish brown. and contrasting white stomatal bands underneath. It 105
Seed scales cyathiform-flabellate, length width is not at all tolerant of droughts and performs best in
at mid-cone 1.52.2 22.5cm; surface smooth, cool, wet conditions but on light, well-drained soils.
puberulent on exposed parts; upper margin entire, It is mostly restricted to collections in botanic gar-
incurved; base pedicellate. Bracts obovate or obcor- dens and arboreta.
date, 11.5cm long, included. Seeds conical-ovoid,
(4)57mm long, light brown; seed wings cuneate,
1012 8mm, light brown, with a purplish tinge. Abies nebrodensis (Lojac.) Mattei, Boll. Reale Orto
Bot. Palermo 7: 64. 1908. Abies pectinata Gilib. var.
Distribution nebrodensis Lojac., Fl. Sicula 2 (2): 401. 19041907;
Abies alba Mill. var. nebrodensis (Lojac.) Svoboda,
Japan: Honshu. Trudy Bot. Inst. Akad. Nauk S.S.S.R., ser. 1, Fl.
TDWG codes: 38 JAP-HN Sist. Vyss. Rast. 13: 60. 1964; Abies alba Mill. subsp.
nebrodensis (Lojac.) Nitz., Lustgrden 1968: 178.
Ecology 1969. Type: not designated. Fig. 12
A species of the high mountain sides and ridges in Abies pectinata Guss., Fl. Sicula Syn. 2: 614. 1844, non
the upper montane and subalpine zones, occurring Gilib. (1792).
commonly between 1000 m and 2800 m a.s.l. (as low
as 750 m in N Honshu). The soils are mostly derived Etymology
from volcanic rock, usually podzolic and slightly
acid or neutral, well drained, and moderately moist The species epithet refers to the Monti Nebrodi, an
(mesic). The climate is cold, with abundant winter alternative name for the Madonie Mountains where
snow and cool, moist summers, the annual precipi- this species was discovered.
tation exceeds 2000mm in the mountains nearest to
the Sea of Japan. Frequent typhoons are a destruc- Vernacular names
tive force reducing the maximum age of trees. Abies
mariesii forms sometimes pure forests near the tree Sicilian fir
line, but is more common in mixed (coniferous) for-
ests with e.g. Abies veitchii, Tsuga diversifolia, Picea Description
jezoensis var. hondoensis and/or undergrowth of
Pinus pumila and Juniperus communis var. nippon- Trees to 1015 m tall in Sicily at present, but may
ica, the latter two especially abundant on ridgetops. grow taller, d.b.h. to 0.40.6 m; trunk monopo-
Common broad-leaved trees are Betula ermanii, dial, straight, columnar, terete; crown (broad)
Sorbus commixta, and Acer spp. In many previously conical (the only living old tree in the wild had its
disturbed areas with deep, fine textured soil, e.g. vol- top broken off). Bark of young trees smooth, light
canic ash, there is a dense cover of small bamboo grey, becoming rough and scaly with age, fissured
(Sasa paniculata and S. nipponica), which excludes at base. Branches of first order spreading horizon-
most other plants (Franklin et al., 1979). tally; branches of second order idem. Branchlets
stout, firm, yellowish green, shiny, turning grey, with limit, Fagus sylvatica are the most common low trees.
prominent ridges between the leaves, glabrous, or Juniperus communis forms dense ground covering
minutely pubescent in the first year; leaf scars circu- carpets and could provide protection for seedlings
lar, with a light central part. Vegetative buds conical, of A. nebrodensis against dehydration and/or grazing
89 46mm, slightly resinous; bud scales triangu- and browsing animals.
lar-ovate, laciniate, protruding, light brown, persist-
ing several years. Leaves spirally arranged, spreading Conservation
laterally in two sets, or the upper leaves covering
shoot and directed forward, the lower leaves pec- Abies nebrodensis is one of the rarest conifers in the
tinate on shaded shoots, leaves on coning shoots world, with only 29 individual plants remaining in
106 assurgent, (1)1.52(2.2) cm long, 23.5mm wide, the wild in 2006. Most of these are small trees grow-
twisted or curved at base, linear-ligulate, flattened, ing on heavily overgrazed scree slopes with clumps
longitudinally grooved above, bright, lustrous green of Fagus sylvatica and Quercus petraea scatterd
on the adaxial (upper) surface, two greenish white among them. Extensive logging since the beginning
bands on the lower surface; apex variable: acute or of the 18th century had brought this species to the
mucronate, on shaded shoots obtuse but not emar- brink of extinction when it was discovered nearly
ginate. Stomata none or a few near the apex above, in a century ago in a nearby village, and more recently
two bands separated by a midrib below. Pollen cones the remaining trees in the valley of Madonna degli
lateral, crowded, 1.52cm long, greenish yellow with Angeli on Mt. Scalone. An extensive ex situ con-
purple microsporophylls. Seed cones lateral, erect, servation programme both locally and abroad is in
short pedunculate, cylindrical, with a conical apex, operation but attempts at re-introduction have to
(7)810(12) cm long, 34cm wide, yellowish green date not been very successful due to harsh summer
when immature, ripening to greenish brown (light conditions and a totally depleted soil. New attempts
reddish brown inside); cone rachis persistent, nar- funded by the EU and providing initial planting
rowly conical, brown. Seed scales cuneate-cyathi- compost and watering in summer may be more
form, length width at mid-cone 22.5 2.83.3cm; successful; protection from direct sunlight, erosion
surface smooth or slightly wrinkled, puberulent on and grazing are also essential. Ultimately the forest
exposed parts; upper margin entire or undulate; ecosystem has to be restored for the survival of this
base pedicellate. Bracts linear-spathulate, 2.53cm relict conifer.
long, exserted and recurved. Seeds conical-oblong, IUCN: CR (D)
68mm long, reddish brown; seed wings cuneate,
oblique, 1015mm long, light brown. Uses
Hinggan fir, Khinghan fir; chou lengshan (Chinese); A natural hybrid between A. sibirica and A. nephro-
Pikhta amurskaya, Pikhta belokoraya (Russian) lepis has been reported from Heilongjiang Province,
China: Abies sibirico-nephrolepis Taken. & Chien.
Description Its leaves are described as being shorter than those
of A. nephrolepis and its young shoots are greyish
Trees to 3035 m tall, d.b.h. to 11.2 m; trunk mono- brown pubescent, while the seed cones are larger.
podial, straight, columnar, terete; crown conical Other character states are intermediate between the
or oval, old trees with densely branched flat tops two parent species. It is reported to be common in
(storks nests). Bark of young trees smooth, light river valleys in the Lesser Hinggan Range. The occur-
grey to greyish brown, in old trees shallowly fis- rence of A. sibirica in this region is not confirmed by
sured, dark greyish brown. Branches of first order other accounts (e.g. Flora of China 4, 1999) and the
spreading horizontally, those near the top ascend- hybrid status of the firs in this area is doubtful.
ing, the lowest drooping; branches of second order
spreading horizontally, dense. Branchlets slender, Distribution
firm, yellowish grey brown, turning to grey; surface
ridged and grooved, with minute pubescence in the Russian Far East: from the Zeya River to the Sikhote
grooves; leaf scars circular. Vegetative buds ovoid or Alin Range; NE China: Manchuria, Shaanxi, south to
conical, broad, 5 4mm, resinous, especially near Shanxi (Wutai Shan); North Korea and South Korea.
apex; bud scales triangular, blunt, brown or pur- TDWG codes: 31 AMU KHA PRM 36 CHM CHN-HB
plish red, appressed, persisting several years. Leaves CHN-SA 38 KOR-NK KOR-SK
spirally arranged, radially around shoot, directed
forward and assurgent, covering the upper part Ecology
of shoot, 12.5(3) cm long, 2mm wide, strongly
twisted at base, linear, flattened, with slightly revo- This is a species of low to medium high mountains,
lute margins, dull, light (grey-)green, with whit- occurring at elevations between 500 m and 700 m
ish green bands below; apex variable: emarginate, a.s.l. in E Siberia at the northern limit of its range,
108
plate 3. Abies nephrolepis. 1. Habit of trees. 2. Foliage branch. 3. Branch with seed cones. 4. Detail of
foliage. 5, 6. Leaves. 7. Seed scale with bract. 8. Seeds.
between 750 m and 2000 m a.s.l. in NE China. This or pyramidal, old trees often flat topped. Bark of
species grows on a variety of well drained mountain young trees smooth, grey, in old trees rough, shal-
soils. The climate is cold, with short, cool and moist lowly fissured, blackish grey-brown. Branches of first
summers and long, cold winters. Most of the annual order spreading, ascending in upper part of crown;
precipitation is snow. It is usually associated with branches of second order spreading horizontally,
other conifers, e.g. Pinus koraiensis and Picea jezoen- assurgent near the top. Branchlets slender, firm, light
sis; also with Pinus pumila and Juniperus sabina var. olive brown or brown, ridged and grooved, with
davurica at higher elevations (maritime provinces of brown pubescence in grooves or glabrous; leaf scars
the Russian Far East); in the interior with Picea obo- circular, light. Vegetative buds ovoid, with pointed
vata, Larix gmelinii, Pinus sibirica or Abies sibirica. apex, 6 5mm, not resinous or resinous; bud
Betula spp. and Sorbus amurensis are common asso- scales ovate, acute, keeled and with laciniate edges, 109
ciated broad-leaved trees. red-brown, persisting several years. Leaves spirally
arranged, pectinate below, the upper leaves pressed
Conservation forward above shoot, on coning shoots assurgent,
(1.5)23(3.5) cm long, 1.52.5mm wide, strongly
IUCN: LC twisted at base, linear, flattened, sometimes with
slightly revolute margins, grooved above, lustrous
Uses dark green above, two whitish green bands below;
apex emarginate, obtuse on coning shoots. Stomata
Hinggan fir is an important timber tree in NE China in two bands separated by a midrib below. Pollen
and Korea. Its wood is used in carpentry and for ply- cones lateral, crowded, pendulous, 12cm long,
wood and veneer. The relatively small size of this tree yellowish. Seed cones lateral, erect, often crowded,
on marginal sites makes exploitation commercially short pedunculate, ovoid-cylindrical, with pointed
unlikely; the better, larger trees come from mixed and often papilliform apex, (10)1216(20) cm
conifer forests at middle elevations in the mountains. long, 45.5(6) cm wide, greenish when immature,
In horticulture, it was introduced to Great Britain ripening to light brown (reddish brown inside); cone
in 1908 from the botanic garden in St. Petersburg, rachis persistent, narrowly conical, dark brown.
Russia. It remains a rarely planted species, which is Seed scales flabellate or cyathiform, length width
susceptible to damage by late spring frosts in coun- at mid-cone 1.82.5 2.74cm; surface smooth,
tries with an Atlantic maritime climate. slightly striated, pubescent on exposed parts; upper
margin entire, often repand, slightly incurved; base
pedicellate. Bracts spathulate-obcordate, with lacin-
Abies nordmanniana (Steven) Spach, Hist. Nat. iate edges and a short or long cusp with midrib,
Vg. Phan. 11: 418. 1841. Pl. 4 2.73.5cm long, exserted and reflexed. Seeds cune-
ate, 1012 78mm, fawn brown, shiny; seed wings
Etymology cuneate, with rounded edge, 1518 15mm, light
purplish brown or rose, turning fawn brown.
This species was named after the botanist A. von
Nordmann, who introduced it to horticulture in Taxonomic notes
1838.
It is now generally accepted that the firs of NW Turkey
Vernacular names are conspecific with Abies nordmanniana, which has
its main distribution in the Caucasus Mountains.
Caucasian fir, Nordmann fir; Pikhta kavkazkaya The disjunct populations of NW Turkey have been
(Russian) known as and are sometimes still accepted as two
distinct species: A. bornmuelleriana (Ulu-Dagh)
Description and A. equi-trojani (Kaz-Dagh), but the morpho-
logical differences between the two are minor and
Trees to 50 m tall, d.b.h. to 1.52 m; trunk monopo- the character states are overlapping, not discrete. It
dial, straight, columnar, terete; crown broad conical seems better to recognize just one subspecies for the
110
plate 4. Abies nordmanniana. 1. Habit of tree. 2. Foliage. 3. Seed cone. 4, 5. Seed scales with bract (4),
with seeds (5). 6. Bract. 7. Leaves.
populations of NW Turkey. If recognized as a dis- Abies nordmanniana (Steven) Spach subsp. nord-
tinct subspecies of A. nordmanniana, the earliest manniana. Pinus nordmanniana Steven, Bull. Soc.
combination made at that rank, A. nordmanniana Imp. Naturalistes Moscou 11: 45, t. 2. 1838. Type:
subsp. equi-trojani, is the valid name. Georgia: Caucasus Mts. (South), [source of the
Kur River], A. von Nordmann s.n. in herb. Steven
Distribution (holotype H).
Distribution Description
114
Mountains from Afghanistan east to Nepal, Leaves 36(7) cm long, on vegetative shoots pec-
Karakoram Range in Pakistan. tinately arranged; apex sharply bifid or emarginate;
TDWG codes: 34 AFG 40 NEP PAK WHM-HP stomata only in two bands on the underside.
WHM-JK WHM-UT
Distribution
Ecology
Mountains from Afghanistan east to Nepal,
Abies pindrow is a species of high mountains, occur- Karakoram Range in Pakistan.
ring between 2000 m and 3300 m a.s.l. (occasion- TDWG codes: 34 AFG 40 NEP PAK WHM-HP
ally as high as 3700 m; Liu, 1971), on alpine lithosols. WHM-JK WHM-UT
The climate is cool, moist monsoon, with abundant
precipitation, but less than in the eastern Himalayas, Conservation
much of it falling as snow. It occurs in pure stands
or in association with Picea smithiana, Pinus walli- IUCN: LC
chiana, Tsuga dumosa and Cedrus deodara; at lower
elevations broad-leaved trees, e.g. Quercus semecar-
pifolia, Q. dilatata, Juglans regia, Aesculus indica, Abies pindrow (Royle ex D. Don) Royle var.
Acer spp., Prunus spp., and Ulmus spp. become more brevifolia Dallim. & A. B. Jacks., Handb. Conif.:
important, replacing the conifers below 1600 m. 126. 1923. Type not designated.
Description Ecology
Trees to 30 m tall, but most trees in nature smaller, Abies pinsapo is a species of the north slopes of high
d.b.h. to 11.5 m; trunk monopodial, straight, colum- mountains, where it occurs between 900 m and
nar and terete in sheltered trees, but often twisted 1800 m a.s.l. (var. pinsapo in S Spain) and 1400 m
and forked; crown in young trees narrowly conical, and 2100 m a.s.l. (the Moroccan variety). It grows on
old trees irregular, open or dense. Bark of young rocky soils derived from dolomitic limestone or ser-
trees smooth, dark grey, in old trees rough and scaly. pentine, with deep drainage. The climate is montane,
Branches of first order long, curved downward, the with a mediterranean influence: dry, warm summers
ones near the top ascending; branches of second alternate with cool, moist winters, with annual pre-
order dense, spreading horizontally and ascending. cipitation around 1000mm. Both varieties occur in
Branchlets stout, very firm, reddish brown or green- pure, scattered stands (very rare in Spain) or mixed
ish brown, turning grey, faintly ridged between the with Cedrus atlantica (Morocco) or Pinus pinaster
leaves, glabrous; leaf scars circular or angular, large, (Spain). Commonly, broad-leaved trees, e.g, Quercus
purplish grey. Vegetative buds ovoid-globose, 56 ilex, Q. lusitanica, and Q. canariensis (Morocco), and
44.5mm, not resinous to very resinous; bud (often sclerophyllous) shrubs, e.g. Ulex balticus,
scales triangular, keeled and free at the apices, light Cistus spp., Pistacia lentiscus, Daphne laureola and
reddish brown or purplish brown. Leaves spirally Berberis hispanica are mixed with scattered A. pin-
arranged, spreading radially and perpendicularly sapo at lower elevations.
from shoots, or more or less pectinate, the upper
leaves often recurved, in shaded shoots the lower Uses
leaves somewhat pectinate, 0.62cm long, 23mm
wide, not or only slightly twisted at base, linear-lig- Spanish fir (including its Moroccan variety) is no
ulate, carinate or slightly flattened, rigid, grey-green longer exploited for its timber. This species is one of
or glaucous green; apex obtuse, acute or acuminate. the most attractive firs in horticulture with its rigid
Stomata above in several rows, below in two bands needles spreading all around the shoots and espe-
separated by a midrib and bordered by broad mar- cially some more glaucous-leaved trees have been
gins. Pollen cones lateral, crowded, 0.50.7cm long, selected as cultivars and are widely planted. While
yellowish with red or violet microsporophylls. Seed most remaining trees at high altitude in southern
cones lateral, erect, short pedunculate, cylindrical, Spain are short and slow growing, Spanish fir can
grow tall in less exposed sites with ample moisture. Park grazing by goats, a pan-Mediterranean men-
For garden purposes slower growing dwarf forms ace, hampers natural regeneration. Erosion of the
are more popular and a few cultivars, some derived already thin and depleted soil adds to the low ability
from witches brooms found in natural populations, to establish new trees naturally. A planting scheme to
meet these requirements. Apparently this species restore depleted populations is being implemented.
can be easily hybridized with a number of other spe- IUCN: EN [B1ab(i, ii, iii)+2ab(i, ii, iii)]
cies, e.g. A. alba, A. cephalonica, A. numidica (its
closest relative), A. nordmanniana and A. pindrow;
some of these crosses have been the parental stock Abies pinsapo Boiss. var. marocana (Trab.) Ceballos
of further cultivars. Many hybrids resulting from & Bolao, Bol. Inst. Nac. Invest. Agron. 1 (2): 18.
116 (controlled or spontaneous) crossings in cultiva- 1928. Abies marocana Trab., Bull. Soc. Bot. France
tion have been given nothospecific names e.g. Abies 53: 154. 1906. Type not designated.
vilmorinii Matf. for the cross between A. pinsapo
and A. cephalonica. Since seed produced from these Abies tazaotana S. Czar ex Villar, Types Sols Afrique
F1 crossings in cultivation is likely to have been fer- N. 1: 80. 1947; Abies pinsapo Boiss. var. tazaotana
tilized by unknown fir pollen, perpetuation of the (S. Czar ex Villar) Pourtet, Ann. Ecole Natl. Eaux
hybrid is only guaranteed by vegetative propagation 9 (1): 100. 1954; Abies pinsapo Boiss. subsp. tazaotana
or by renewed controlled crossing of the parent spe- (S. Czar ex Villar) R. Govaerts, World Checklist
cies. Such plants should receive cultivar names, not Seed Pl. 1 (1): 6. 1995.
taxon names.
Description
2 varieties are recognized:
Leaves more ore less pectinately arranged, especially
on lower branches, acute to acuminate; resin canals
Abies pinsapo Boiss. var. pinsapo. Type: Spain: marginal. Seed cones 1018cm long, 3.55cm wide.
Malaga, Sierra Bermeja, N of Estepona [Sierra de
la Nieve], E. Boissier s.n. (holotype not located, Distribution
isotype K).
N Morocco (Rif Mts.).
Description TDWG codes: 20 MOR-MO
Distribution
Abies procera Rehd., Rhodora 42: 522. 1940. Type:
S Spain: Prov. Mlaga, Cdiz. USA: Washington, Columbia River, D. Douglas s.n.
TDWG codes: 12 SPA-SP (holotype not located, isotype K). Fig. 16
Conservation Etymology
Abies pinsapo var. pinsapo is restricted to a small The species epithet means slender or tall and refers
number of disjunct populations, the largest one situ- to the shape of the trees in dense forest stands.
ated within the Sierra de las Nieves National Park.
Fires, some started by tourists, are a hazard threat- Vernacular names
ening especially some of the smaller, isolated stands
with destruction. Within and without the National Noble fir
Description Distribution
Trees to 8090 m tall, d.b.h. to 2.53.5 m; trunk USA: NW California, Oregon, Washington (Cascade
monopodial, straight, columnar, terete, often bare of Range, parts of Coast Range).
branches to a considerable height and very regular; TDWG codes: 73 ORE WAS 76 CAL
crown narrowly pyramidal or more conical, in for-
est stands often only or less of total tree height. Ecology
Bark of young trees smooth, thin, with resin blisters,
light grey with a purplish tinge, in old trees rather A magnificent tree, occurring from the foothills of
smooth, except on lower half of trunk where it mountains in W Washington to high mountain sides
breaks into irregular plates, greyish brown. Branches in Oregon, between 60 m and 2700 m a.s.l. It is most 117
of first order spreading horizontally, the lower pen- abundant in the mountains of the Cascade Range, on
dulous, but often contorted in old trees; branches of a variety of mountain soils with ample moisture avail-
second order spreading horizontally and ascending able to the vegetation. The climate is cool temperate,
near the ends of main branches. Branchlets slender, with short summers and snowy winters, the annual
firm, reddish brown in the first year, soon purplish precipitation ranging from 1750mm to 2600mm,
brown, faintly ridged and grooved, pubescent, but much of it as snow. It mainly grows in the Canadian
almost hidden by leaves; leaf scars oval or obovate. Life Zone, but also in the lower Transition Zone,
Vegetative buds ovoid-globose, hidden by leaves, where it can be associated with several other conifers,
3 2mm, slightly resinous; bud scales ovate, dark e.g. Tsuga heterophylla, Picea sitchensis and Thuja
purplish red, persisting several years. Leaves spirally plicata near the coast, Pseudotsuga menziesii, Abies
arranged, strongly assurgent on all shoots, the lower grandis, Pinus spp. in much of its range, and Abies
leaves spreading and S-curved, 12.5(3.5) cm long, lasiocarpa, A. amabilis, Tsuga mertensiana, Picea
those midway of a years shoot longest, 1.52mm engelmannii, Larix occidentalis at higher elevations.
wide, curved at base, basi-falcate or falcate-linear, Common shrubs are Rhododendron spp., Vaccinium
flattened or carinate, glaucous green above, glaucous spp. and Ribes spp. Abies procera can be dominant,
white bands below; apex obtuse (rarely notched), but occurs rarely in pure stands (Fowells, 1965).
acute on coning shoots. Stomata in several to many
rows above, in two narrow bands separated by a mid- Conservation
rib below. Pollen cones lateral, pedunculate, solitary
or a few together in leaf axils, 1.52.5cm long, yellow IUCN: LC
with red microsporophylls. Seed cones lateral, erect,
short pedunculate, cylindric-conical, with obtuse or Uses
truncate apex, 1520cm long (30cm on cultivated
trees), 58cm wide, green and often tinged with red Noble fir attains large dimensions and grows an
when immature, maturing to greenish grey, with yel- extremely straight bole under favourable site con-
lowish bracts, ripening to light grey brown with light, ditions. Almost pure natural stands can yield large
orange brown bracts; cone rachis persistent, nar- volumes of timber per ha. Its wood is of higher qual-
rowly conical, purplish brown. Seed scales cuneate- ity than that of other firs in North America due to
flabellate or cyathiform, length width at mid-cone greater strength and indeed its size. Besides general
2.53 2.53.5cm; surface smooth, pubescent to hir- construction and carpentry applications, special
sute, with yellowish grey hairs; upper margin entire, uses have been propellors of airplanes and ladders,
strongly incurved; base long pedicellate. Bracts now mostly replaced by various metals. The odour-
spathulate-obcordate, with laciniate upper margins less, white wood is excellent for making boxes. Young
and a long, curved cusp, 34cm long, exserted, trees make attractive Christmas trees with their
reflexed, covering much of the exposed parts of the dense, upturned glaucous leaves. In amenity planting
seed scales. Seeds cuneate-oblong, 1215mm long, and horticulture this fir is one of the more popular
pale reddish brown; seed wings obovate or cuneate- and commonly used species and several cultivars
oblong, ca. 20 15mm, straw coloured. are known. It was introduced to England by David
Douglas in 1830. It has the largest seed cones of all pedunculate, ovoid or ovoid-oblong, with obtuse or
species, with attractive, exserted yellowish bracts. It papilliform, sometimes umbilicate apex, 58(9) cm
is unsuitable in climates with summer droughts or long, 2.53.5cm wide, bluish purple when immature,
less than a good supply of rain spread evenly in the maturing to blue-grey, ripening to grey-brown; cone
year as its bark tends to split open during dry spells. rachis persistent, thick, cylindro-conical to fusiform,
purplish brown or light brown. Seed scales broadly
flabellate to cyathiform, length width at mid-cone
Abies recurvata Mast., J. Linn. Soc., Bot. 37: 423. 1.52 22.5cm; surface smooth, puberulous; upper
1906. margin thin, entire, erose or irregularly fissured;
base pedicellate. Bracts spathulate, with a short cusp,
118 Etymology 11.5cm long, entirely included, or occasionally the
cusps exserted. Seeds cuneate, 58 34mm, light
The species epithet refers to the often recurved or brown; seed wings broadly cuneate, 610 56mm,
reflexed leaves (needles) on the shoots. violet-blue, turning brown.
Min fir; min kiang lien sha, zi guo leng shan China: SW Gansu, Sichuan, NW Yunnan, SE Xizang
(Chinese) [Tibet].
TDWG codes: 36 CHC-SC CHC-YN CHN-GS CHT
Description
Ecology
Trees to 4060 m tall, d.b.h. to 1.52.5 m; trunk
monopodial, straight, columnar, terete; crown coni- Min fir (both varieties) is a high mountain species
cal or pyramidal, flat topped in old trees. Bark of of SW China, occurring between 2300 m and 3600
young trees smooth, soon with papery flakes, grey or m a.s.l. or even higher. It grows usually on grey-
pink brown, in old trees rough, flaky, breaking into brown mountain podzols. The climate is cold, moist,
small scales, dark grey brown. Branches of first order with annual precipitation between 700mm and
relatively short, horizontally spreading; branches of 1000mm. Both varieties are usually constituents of a
second order similar. Branchlets slender, firm, yel- mixed coniferous forest type, with among other spe-
lowish brown, light brown or pink-brown, later grey, cies A. squamata, Picea likiangensis var. rubescens,
smooth or slightly pubescent, shining and glabrous, P. asperata, and Larix potaninii; Picea purpurea and
ridged between the leaves; leaf scars circular, light Abies fargesii var. faxoniana are mainly found with
grey. Vegetative buds ovoid or broad conical, 58 the typical variety, and A. fabri with var. ernestii.
46mm, resinous, but resin soon disappearing; bud Betula albosinensis is the only common broad-leaved
scales triangular, keeled, light brown, persistent for tree at higher elevations, but lower down the slopes
several years. Leaves spirally arranged, spreading other genera, e.g. Acer, Populus, but also different
radially, more or less pectinate below, strongly assur- conifer species, e.g. Tsuga chinensis, Picea brachytyla
gent to recurved above, especially on cone bearing var. complanata and Pinus armandii become more
shoots and on leading shoots higher in the crown, abundant.
(1.2)1.53.5cm long, 1.92.5 mm wide, twisted
or recurved at base, linear, but the shortest leaves Uses
oblanceolate, flattened, longitudinally grooved
above, glossy green above, 2 greenish white bands A timber tree in western China, heavily exploited
below; apex emarginate, obtuse or acute. Stomata in until recently when the Chinese government finally
two bands separated by a midrib below, occasionally decided to preserve its remaining old growth forests
a few near apex above. Pollen cones lateral, pendant, in the western provinces. Its timber was used mainly
11.5cm long, yellow, with reddish microsporo- for construction and carpentry work. The type col-
phylls. Seed cones lateral, erect, often clustered, short lection (of var. recurvata) was collected by Ernest
H. Wilson on his first expedition to western China Abies recurvata Mast. var. ernestii (Rehd.) C. T.
in 1903; the species was introduced to horticulture Kuan, Notes Roy. Bot. Gard. Edinburgh 41: 536.
in the USA and UK from seed collected by him on 1984. Abies ernestii Rehd., J. Arnold Arbor. 20: 85.
subsequent journeys to the Min River drainage. As 1939; Abies chensiensis Tiegh. var. ernestii (Rehd.)
with most Chinese species in Abies, it remains a den- T. S. Liu, Monogr. Gen. Abies: 135. 1971. Type:
drological collectors item and has not entered the China: Sichuan, Daxue Shan, Kangding, [NE of
common gardening trade. A main reason for this Tachien-lu, Tapao-shan], E. H. Wilson 2090
is undoubtedly the unavailability of seed from its (holotype A). Fig. 17
country of origin for a long period after the efforts
of the early 20th centurys plant collectors came to an Description
end. Renewed collecting, made possible in the last 119
few decades in partnership with Chinese botanists, Buds ca. 8 6mm; shoots yellowish brown, faintly
has been undertaken under more restricting condi- pubescent. Leaves 23.5cm long, more or less
tions and the results have largely remained within straight; apex emarginate on vegetative shoots; resin
the confines of major botanic gardens. Even if trees ducts marginal. Seed cones often with an umbilicate
in cultivation produce viable seed, unless they are apex and fusiform rachis.
grown in complete isolation from other species
of Abies, that seed is likely to produce plants with Taxonomic notes
a mixture of genes from almost any of those other
species. In Flora of China 4: 51 (1999) this variety is treated as
a distinct species, while within A. recurvata no infra-
2 varieties are recognized: specific taxa are recognized.
Distribution
Abies recurvata Mast. var. recurvata. Type: China:
Sichuan, Min River, Songpan, [south of Sung Pan, China: SW Gansu, W Sichuan, NW Yunnan, SE
Min Valley], E. H. Wilson 3021 (holotype K). Xizang [Tibet].
TDWG codes: 36 CHC-SC CHC-YN CHN-GS CHT
Description
Conservation
Buds ca. 5 4mm; shoots usually light brown or
pink-brown, glabrous. Leaves (1.2)1,53cm long, IUCN: VU (A2cd)
often recurved at base; apex obtuse or acute; resin
ducts medial. Seed cones with obtuse or papilliform
apex; rachis usually cylindro-conical. Abies religiosa (Kunth) Schltdl. & Cham., Linnaea
5: 77. 1830. Pinus religiosa Kunth, in Humboldt et
Distribution al., Nov. Gen. Sp. Pl. 2 (5): 5. 1817. Type: Mexico:
Guerrero, F. W. H. A. von Humboldt & A. J. A.
China: SW Gansu, N Sichuan. Bonpland s.n. (holotype P).
TDWG codes: 36 CHC-SC CHN-GS
Abies colimensis Rushforth & Narave, Notes Roy.
Conservation Bot. Gard. Edinburgh 46: 105. 1989.
Sakhalin fir; akatodo, todo-matsu (Japanese); Pikhta Japan: Hokkaido; Russian Far East: southern Kuril
sakhalinskaya (Russian) Islands, Sakhalin.
TDWG codes: 31 KAM KUR SAK 38 JAP-HK
Description
Ecology
Trees to 2530 m tall, d.b.h. to 0.81 m; trunk mono-
podial, straight, columnar, terete; crown broad pyra- Sakhalin fir and its varieties occur from near sea
midal, often flat topped in old trees; dead branches level on the coast to an elevation of 1650 m a.s.l. in
remain on the bole. Bark of young trees smooth, the mountains. The soils are well drained but moist
with resin blisters, grey with a brown tinge, of old throughout the year, due to abundant precipitation
trees in lower part of the bole breaking into irregu- in a cool to cold, maritime climate. In the north of
lar plates, grey brown. Branches of first order long, its range the species is more common at elevations
slender, mostly ascending, the lower ones spread- between 800 m and 1100 m, where it is mixed with
ing horizontally; branches of second order simi- Picea jezoensis, P. glehnii, Larix gmelinii var. japonica
lar. Branchlets slender, firm, red-brown or brown, or Pinus pumila at the highest limit of trees. At lower
sometimes grey-brown, with faint ridges and shal- elevations pure stands occur, below 800 m broad-
low grooves, minutely pubescent in grooves with leaved-trees, e.g. Betula ermanii, Acer spp., Quercus
red-brown hairs; leaf scars circular, small. Vegetative mongolica var. grossesserata, Castanea crenata,
buds ovoid, small, very resinous; bud scales red- Kalopanax septemlobus, and Magnolia hypoleuca
dish brown, persisting several years. Leaves spirally become more abundant.
arranged, spreading more or less radially, the upper
leaves directed forward, covering shoot, on coning Uses
shoots slightly assurgent, 1.23.5cm long, 11.2mm
wide, (strongly) twisted or curved at base, linear, This species is mainly logged for the manufacture
grooved above, flattened, glossy dark green above, of wood pulp used in the paper industry; its timber
two greenish white bands below; apex emarginate is of low quality for construction and carpentry. As
or obtuse. Stomata none or a few near apex above, an amenity tree it is little used outside the cool to
in two narrow bands separated by a midrib below. cold maritime climate of northern Japan and the
Pollen cones lateral, densely clustered on the under- Russian Far East. It is in cultivation in botanic gar-
side of shoots, 1cm long, yellowish, with red micro- dens and arboreta in Russia, northern Europe and
sporophylls. Seed cones lateral, erect, often crowded, New England, USA, but rarely survives to maturity
almost sessile, ellipsoid-cylindrical, with obtuse or in countries with mild winters, where it will not go
acutish apex, 58cm long, 23cm wide, light or dark into prolonged winter dormancy and is susceptible
purple with greenish or reddish bracts when imma- to spring frosts.
4 varieties are recognized, one tentatively: Taxonomic notes
Abies sachalinensis (F. Schmidt) Mast. var. corticosa Russian Far East: Kamchatka (Kronotzky Bay) [pos-
Tatew., Trans. Sapporo Nat. Hist. Soc. 45: 70. 1935; sibly introduced].
Abies sachalinensis (F. Schmidt) Mast. f. corticosa TDWG codes: 31 KAM
(Tatew.) Hayashi, Taxon. Phytogr. Japon. Conif.: 26.
1960. Ecology
IUCN: LC Conservation
IUCN: DD
Abies sachalinensis (F. Schmidt) Mast. var.
gracilis (Kom.) Farjon, Pinaceae (Regnum Veg.
121): 83. 1990. Abies gracilis Kom., Trudy Imp. Abies sachalinensis (F. Schmidt) Mast. var. may-
S.-Peterburgsk. Bot. Sada 20: 203. 1901; Abies riana Miyabe & Kud, Trans. Sapporo Nat. Hist.
sibirica Ledeb. var. gracilis (Kom.) Patschke, Bot. Soc. 7: 131. 1919. Abies mayriana (Miyabe & Kud)
Jahrb. Syst. 48: 684. 1913. Type not designated. Miyabe & Kud, Icon. Ess. Forest Trees Hokkaido 1:
9, t. 3, 4. 1920. Type not designated.
Description
Description
Bark grey. Leaves with marginal resin ducts. Seed
cones purple when immature; bracts exserted, turn- Bark smooth, whitish grey. Leaves with medial resin
ing brown at maturity. ducts. Seed cones light purplish, with greenish,
exserted bracts turning light brown at maturity.
Distribution of first order short, slender, spreading horizontally,
the lower pendant; branches of second order dense,
Japan: Hokkaido; Russian Far East: Sakhalin. spreading out laterally or assurgent. Branchlets slen-
TDWG codes: 31 SAK 38 JAP-HK der, firm, pale yellowish grey to fawn brown or yellow-
ish brown, soon grey, smooth or only faintly ridged
Conservation or more prominently ridged and grooved, with short
pubescence, soon glabrous; leaf scars small, circular.
IUCN: LC Vegetative buds globose, 23mm long, more or less
resinous; bud scales reddish or yellowish brown,
persisting several years. Leaves spirally arranged, the
Abies sachalinensis (F. Schmidt) Mast. var. nemo- upper leaves covering shoot or assurgent, the lower 123
rensis Mayr, Monogr. Abiet. Japan. Reich.: 42, t. 3, f. leaves pectinate in two lateral planes, on coning
6. 1890. Abies nemorensis (Mayr) Miyabe & Kud, shoots all leaves assurgent, (1)1.53cm long, 1.3
in Miyabe & Miyake, Fl. Saghalin: 745. 1915. Type 1.6mm wide, twisted or curved at base, linear, flat-
not designated. tened, light green, with 2 greyish green bands below;
apex emarginate, obtuse, or acute with a callous tip
Description especially on coning shoots. Stomata occasionally a
few near apex above, in two bands separated by a
Bark grey. Leaves with medial resin ducts. Bracts of midrib below. Pollen cones lateral, crowded on the
seed cones included or only the small cusps exserted. underside of shoots, 1.5cm long, yellow, with red
microsporophylls. Seed cones lateral, erect, nearly
Distribution sessile, ovoid-cylindrical, with obtuse apex, 57.5cm
long, 2.53.5cm wide, violet blue when immature,
Japan: Hokkaido; Russian Far East: Sakhalin. maturing to bluish brown, or yellowish brown,
TDWG codes: 31 SAK 38 JAP-HK becoming (light) cinnamon or greenish brown when
ripe; cone rachis persistent, narrowly conical, dark
Conservation brownish purple. Seed scales broad flabellate, upper
scales more cuneate, length width at mid-cone 1.5
IUCN: DD 1.7 22.2cm; surface smooth, puberulent; upper
margin entire or serrulate-erose; base pedicellate.
Bracts short, rounded, with a tiny cusp, 0.8cm long,
Abies sibirica Ledeb., Fl. Altaica 4: 202. 1833. entirely included. Seeds cuneate-oblong, 56mm
long, brown; seed wings cuneate-oblong, 1012mm
Etymology long, light brown.
Description IUCN: LC
Across N Russia, from Archangelsk eastward to the Abies webbiana (Wall. ex D. Don) Lindl. var. brevi-
Amur River, southward to the mountains along the folia A. Henry, in Elwes & Henry, Trees Gr. Brit.
Sino-Russian border. Ireland 4: 751. 1909; Abies spectabilis (D. Don) Spach
TDWG codes: 14 RUE RUN 30 ALT BRY CTA IRK var. brevifolia (A. Henry) Rehd., J. Arnold Arbor. 1:
KRA TVA WSB YAK 31 AMU KHA 36 CHX 54. 1919.
Abies spectabilis (D. Don) Spach var. langtangensis mid-cone 1.52.5 33.5cm; surface smooth, slightly
Silba, Phytologia 68: 22. 1990. striated, puberulent on exposed parts; upper margin
entire, undulate or incurved; base pedicellate, lat-
Etymology erally auriculate. Bracts spathulate, with rounded,
serrulate margin and small cusp, 1.52cm long, usu-
The species epithet means splendid or superb. ally included, or slightly exserted near base of cone.
Seeds cuneate, 10 5mm, brown; seed wings cune-
Vernacular names ate-oblong, 12 5mm, brown-violet.
Description Ecology
Trees to 40 m tall, d.b.h. to 2 m; trunk monopodial, A subalpine species of the high mountains of west-
straight, columnar, terete; crown broad or narrow, ern China, where it occurs between 3500 m and 4500
conical. Bark of young trees smooth, purplish or m a.s.l. [30004700 m according to Liu (1971)] mak-
pink brown, exfoliating like Betula, the bark in older ing it one of the highest reaching mountain trees
trees remaining shaggy from flakes of papery bark, in the world. The soils are commonly grey-brown
breaking into rough and hard plates, orange-brown mountain podzols or lithosols. The climate is cold,
(when freshly exposed), blackish grey at base of relatively dry (arid in E Xizang), but usually per-
trunk. Branches of first order long, spreading hori- petual snow at higher elevations provides sufficient
zontally, the lower pendant; branches of second order moisture throughout the year. It is a constituent of
spreading horizontally, assurgent near the top of the mixed coniferous high altitude forests, with among
tree. Branchlets stout, firm, dark reddish or purplish other species Abies recurvata, A. fargesii var. faxoni-
brown, shining, ridged and grooved, usually gla- ana, Picea likiangensis var. rubescens, P. asperata, P.
brous or a few hairs in the grooves; leaf scars circu- linzhiensis (in E Xizang), Larix potaninii and possibly
lar. Vegetative buds ovoid-globular, thickly covered also Tsuga forrestii. There are very few broad-leaved
with white, soon eroding resin; bud scales triangu- trees at these high elevations, Betula albosinensis and
lar-ovate, keeled, reddish brown, persisting several B. utilis var. prattii being the most common.
Conservation first order slender, relatively short, spreading hori-
zontally, ascending near the top; branches of second
At these high altitudes forests form isolated patches order similar. Branchlets slender, firm, young shoots
on favourable sites, surrounded by treeless subal- green or light brown, yellowish grey in second year;
pine vegetation. Direct exploitation of the timber in surface smooth, in later years with shallow grooves,
these forest remnants is easily unsustainable due to densely yellowish pubescent, soon glabrous; leaf
very slow growth and past exploitation has led to a scars circular, yellowish brown. Vegetative buds
decline of this and other conifer tree species in these ovoid globular, 3 3mm, resinous; bud scales trian-
forests. gular, with erose margins, reddish or purplish brown.
IUCN: VU (A2d) Leaves spirally arranged, radially spreading, the
lower leaves more or less pectinate, the upper leaves 127
Uses directed forward, covering shoot, on coning shoots
slightly assurgent, (0.5)1.53(3.8) cm long, 1.5
Flaky fir is a potential timber tree but its occur- 2.2mm wide, widest near the emarginate or truncate
rence at extremely high altitudes in inaccessible apex, linear-ligulate or falcate, grooved above, flat-
places prevents it from being exploited commer- tened, dark green above, two whitish bands below.
cially. Ernest Wilson collected this fir with its pecu- Stomata in two bands separated by a midrib below.
liar bark in June 1904 in the Daxue Shan of western Pollen cones lateral, axillary, pendant, 11.5cm long,
Sichuan, China, when on a plant hunting expedition yellowish, with red microsporophylls. Seed cones
for Veitch & Sons in England. Although it was suc- lateral, erect, often crowded, short pedunculate or
cessfully introduced in Europe and North America, almost sessile, cylindrical to ellipsoid (often irregu-
it has remained rare in cultivation, restricted to a few lar), with obtuse or papilliform apex, (3)4.57.5(8)
collections in botanic gardens and arboreta, where cm long, (1.5)22.5(3) cm wide, dark bluish pur-
it tends to be a slow grower. Its unusual bark has an ple (rarely green or olive-green) when immature,
attraction to dendrologists, but unless renewed seed ripening to blackish brown or brown; cone rachis
collecting from wild sources can be resumed, this persistent, narrowly conical, purplish brown. Seed
species may gradually disappear from horticulture. scales narrowly reniform or almost crescent shaped,
length width at mid-cone 0.81 1.41.6cm; sur-
face smooth, (silvery) puberulent on exposed parts;
Abies veitchii Lindl., Gard. Chron. 1861: 23. 1861. upper margin entire, incurved; base pedicellate.
Bracts obcordate, 11.2cm long, slightly exserted or
Etymology only the cusps exserted, straight or recurved. Seeds
cuneate, 56 3mm, greyish black tinged with
This species has been named after John Gould green; seed wings broadly cuneate, 3 5mm, colour
Veitch, founder of Veitch Nurseries, who discov- as seeds or blackish purple.
ered it in 1861 on Mt. Fuji.
Distribution
Vernacular names
Japan: Honshu, Shikoku.
Veitchs fir; shira-biso (Japanese) TDWG codes: 38 JAP-HN JAP-SH
Description Ecology
Trees to 2530 m tall, d.b.h. to 0.81 m; trunk mono- The typical variety of this species grows on high
podial, straight, columnar, terete; crown (narrowly) mountains at elevations between 1200 m and 2800
pyramidal. Bark of young trees smooth, with promi- m a.s.l. [reported from as low as 1050 m (Wilson,
nent resin blisters, (light) greenish grey, lower part 1916)]. The soils are usually of volcanic origin,
of trunk becoming scaly and dark grey. Branches of podzolic and well drained. The climate is cool and
wet, with annual precipitation between 1000mm Conservation
and 2500mm, and with cold, snowy winters; fre-
quent typhoons cause destruction of the forest in IUCN: LC
most places before it reaches an age of 250 to 300
years (Franklin et al., 1979). This variety is usually
mixed with other conifers, e.g. Abies mariesii, Picea Abies veitchii Lindl. var. sikokiana (Nakai) Kusaka,
jezoensis subsp. hondoensis, Larix kaempferi, Thuja Conif. Japon. Ill., ed. 2: 212. 1954. Abies sikokiana
standishii, Pinus parviflora, at the highest elevations Nakai, Bot. Mag. (Tokyo) 42: 452. 1928. Type not
Pinus pumila, and the ubiquitous Tsuga diversifolia. designated.
The most common broad-leaved trees are Betula
128 ermanii, Sorbus commixta, Prunus nipponica, and Description
Acer spp. at lower elevations, and Betula corylifolia
near the tree limit. Leaves (0.5)0.82cm long, 2mm wide. Seed cones
ellipsoid-cylindrical, 34cm long, 1.52mm wide;
Uses only the bract cusps are exserted.
2 varieties are recognized: Abies vejarii Martnez, Anales Inst. Biol. Univ. Nac.
Mxico 13 (2): 629. 1942.
NE Mexico: Sierra Madre Oriental in Coahuila, Rushforth (1987) has rejected alliance of this variety
Nuevo Len and Tamaulipas. with A. vejarii and placed this taxon close to Abies
TDWG codes: 79 MXE-CO MXE-NL MXE-TA durangensis.
Conservation Distribution
Distribution Conservation
Etymology
Abies vejarii Martnez var. mexicana (Martnez)
T. S. Liu, Monogr. Gen. Abies: 261. 1971. Abies The epithet refers to the mountain where this species
mexicana Martnez, Anales Inst. Biol. Univ. Nac. was discovered.
Mxico 13 (2): 626. 1942; Abies vejarii Martnez
subsp. mexicana (Martnez) Farjon, Pinaceae Vernacular names
(Regnum Veg. 121): 103. 1990. Type: Mexico: Nuevo
Len, Sierra de Santa Catarina, M. Martnez s.n., Yuanbaoshan fir; Yuanbaoshan lengshan (Chinese)
1939 (holotype MEXU).
Description
Description
Trees to 25 m tall, d.b.h. to 0.6 m (or more); trunk
Seed cones to 10cm long; bracts with a narrow, monopodial, straight, columnar, terete; crown prob-
straight cusp, entirely included. ably like A. forrestii (not described). Bark smooth
at first, becoming ridged and grooved, divided into
small plates in old trees. Branches of first order long,
spreading horizontally; branches of second order
spreading and ascending. Branchlets slender, firm,
yellowish brown or light brown, becoming light or Ecology
dark brown, glabrous; leaf scars circular. Vegetative
buds ovoid-conical, very resinous; bud scales red- The highest mountains in Guangxi, like Yuanbao
dish brown, persisting several years. Leaves spirally Shan, have a very cool, wet climate, with annual pre-
arranged, densely set, subradially spreading above cipitation exceeding 2000mm. The summers are cool
shoot, the lower leaves spreading laterally, the upper and cloudy, the winters last 45 months and bring
leaves shortest, on coning shoots assurgent or erect, abundant snow from December through March. This
12.7cm long (leaves of young trees longer: 33.8cm, species occurs in mixed deciduous-coniferous forest
usually more pectinately arranged in two rows of with other conifers (e.g. Tsuga chinensis) and broad-
equal length), longest in the middle portion of shoot, leaved trees dominated by members of the Fagaceae;
1.82.5mm wide, twisted or curved at base, linear the Abies trees are very scattered. 131
or ligulate-linear, flattened; margins (in sicco) revo-
lute, with a median groove above, dark green above, Conservation
two very white bands below; apex obtuse to slightly
emarginate. Stomata absent above, in two broad This species has an extremely limited distribution,
bands separated by a midrib below. Pollen cones distant from other species, and is only known from
solitary, axillary, on short branches, oblong cylindri- one small area in Guangxi Province.
cal, 11.5cm long, yellow. Seed cones lateral, erect, IUCN: CR [B1ab (v) + B2ab (v)]
short pedunculate to almost sessile, broad ovoid-
cylindric, short, with slightly narrowed, obtuse apex, Uses
89cm long, 4.55cm wide, green or yellowish green
when immature, maturing to light yellowish brown, No uses have been recorded of this species.
becoming pale (yellowish) brown when ripe; cone
rachis persistent, fusiform or conical, brown. Seed
scales cuneate-flabellate, length width at mid cone Abies ziyuanensis L. K. Fu & S. L. Mo, Acta
2 2.2cm; surface smooth, with thickened central Phytotax. Sin. 18 (2): 208. 1980. Abies fabri (Mast.)
portion of apical part, exposed part densely greyish Craib var. ziyuanensis (L. K. Fu & S. L. Mo) Silba,
white puberulent; upper margin entire, rounded to Phytologia 68: 14. 1990; Abies beshanzuensis M.
almost truncate, slightly incurved; lower margins H. Wu var. ziyuanensis (L. K. Fu & S. L. Mo) L. K.
auriculate; base short pedicellate. Bracts large, broad Fu & Nan Li, Novon 7 (3): 261. 1997. Type: China:
oblong, widest in the middle (9mm), with rounded, Guangxi, Ziyuan Xian, Yinzhulao Shan, Y. J. L
erose-denticulate upper margins and a small cusp, 78001 (holotype PE).
2.52.8cm long, of the same colour as the seed
scales, apical part 67mm wide, reflexed, distinctly Abies dayuanensis Q. X. Liu, Bull. Bot. Res. North-
exserted. Seeds cuneate-oblong or obovate-oblong, East. Forest. Inst. 8 (3): 85. 1988.
10 4mm, resinous, dark red brown; seed wings
cuneate-dolabriform, with a truncate apex, slightly Etymology
longer than the seeds (911 810mm), shining
light brown. The epithet refers to the name of the municipal-
ity (Ziyuan Xian) in which the type specimen was
Taxonomic notes collected.
Distribution Description
China: N Guangxi (Rongshui Xian, Yuanbao Shan). Trees to 30 m tall, d.b.h. to 0.60.9 m; trunk monopo-
TDWG codes: 36 CHS-GX dial, straight, columnar, terete; crown broad, conical
or flat topped in old trees. Bark of young trees smooth, Momi, subsection Holophyllae, as its close affinity to
grey, in old trees shallowly ridged and grooved, bro- A. chensiensis seemed more likely on morphological
ken into small plates on lower part of trunk. Branches grounds. In Flora of China 4: 50 (1999) A. ziyuanen-
of first order long, spreading horizontally, ascending sis has been reduced to a variety of A. beshanzuensis.
towards the top of the tree; branches of second order If this taxonomy is accepted, A. beshanzuensis would
spreading or ascending. Branchlets thick, stout, at first no longer be considered an extremely rare species,
light yellowish brown, in the third year greyish brown; but merely a variety of a still rare, but more wide-
surface ridged and grooved between the leaves, gla- spread species. A recent phylogenetic study (Xiang
brous or with short hairs in the grooves; leaf scars et al., 2009) based on nuclear DNA sequence data
circular. Vegetative buds cylindric or ovoid-oblong, (ITS regions) sampled A. ziyuanensis amongst 48
132 with acutish apex, covered with a thin layer of white species. This species appeared to be closely related to
resin; bud scales triangular, dorsally keeled, light yel- A. homolepis and also to A. chensiensis.
lowish brown. Leaves spirally arranged, spreading lat-
erally in two overlapping sets, of unequal length, the Distribution
longest near base of shoot, on coning shoots upper
leaves assurgent, all leaves on vegetative shoots of China: NE Guangxi (Rongshui Xian, Yuanbao
young trees pectinate, widely spaced, 24.8cm long, Shan), SW Hunan (Ziyuan Xian, Xingni, Chenbu),
33.5mm wide, twisted or curved at base, linear, SW Jiangxi (Jinggang Shan).
straight or curved, flattened, with margins (in sicco) TDWG codes: 36 CHS-GX CHS-HN CHS-JX
slightly recurved, dark green above, two greenish
white bands below; apex obtuse or slightly emargin- Ecology
ate. Stomata absent on the adaxial (upper) surface, in
two bands divided by a midrib below. Pollen cones Abies ziyuanensis is a rare fir occurring on the high-
lateral, in leaf axils, ca. 2cm long, yellowish, with red est mountains in Jiangxi, Guangxi and on the border
microsporophylls. Seed cones lateral, with 0.51cm with Hunan, in a narrow belt between 1650 m and
long peduncles, oblong-cylindric or elliptical, with 1750 m a.s.l. These mountains have a cool, very wet
obtuse apex, 1011cm long, 4.24.5cm wide, green- climate, with a mean annual temperature between
ish or yellowish green when immature, maturing to 9.212 C, and a winter period of 45 months
dark greenish brown, becoming dark brown when (November-March) in which the mean tempera-
ripe; cone rachis persistent, narrowly conical. Seed ture is between -3 to -5 C (min. -10). The weather
scales broad cuneate-flabellate, length width at mid is usually cloudy, with much fog, the annual pre-
cone 2.32.5 33.3cm; surface smooth, dark brown cipitation is 21002400mm and snow lasts from
when ripe, sparingly puberulent; upper margin entire, December through March. Abies ziyuanensis occurs,
rounded, not incurved, light brownish green; lateral together with other conifers, scattered in a mixed
margins finely toothed, auriculate near the pedicellate forest dominated by deciduous broad-leaved trees.
base. Bracts oblong-spathulate, near apex 910mm Above 1700 m on Yuanbao Shan it is replaced by
wide, near base 3mm, 2.12.3cm long including the A. yuanbaoshanensis.
small cusp, included, or slightly exserted near base
of cone and recurved; apical margin with fine teeth. Conservation
Seeds cuneate-oblong, ca. 10 4mm, purplish grey
with dark resin; seed wings cuneate-dolabriform, with This species is only known from less than five locali-
rounded or slightly truncate apex, 1315 1214mm, ties, some in close proximity. As it occurs lower on
light purplish grey, with dark spots, lustrous. the mountains, it has been logged for local use of
timber. Present threats are landslides and overgraz-
Taxonomic notes ing by sheep and cattle.
IUCN: CR [B1ab (iii) + C2a (i)]
According to Fu, Lu & Mo (1980), this species is
closely related to A. beshanzuensis M. H. Wu, another Uses
novelty described around that time. Farjon &
Rushforth (1989) classified A. ziyuanensis in section No current uses have been recorded of this fir.
Acmopyle Pilg., in Engler, Pflanzenreich 4 (5, 18): 117. 1903. Type: Acmopyle pancheri
(Brongn. & Gris) Pilg. (Podocarpaceae).
Greek: akme = highest point, tip; pyle = opening (gate); Acmopyle pancheri (Brongn. & Gris) Pilg., in
referring to the erect position of the mature seed. Engler, Pflanzenr. IV.5 [18]: 117. 1903. Dacrydium
pancheri Brongn. & Gris, Bull. Soc. Bot. France 16:
Description 330. 1869; Nageia pancheri (Brongn. & Gris) Kuntze,
Revis. Gen. Pl. 2: 800. 1891. Type: New Caledonia:
Dioecious (or sometimes monoecious? ), small Grande Terre, Province Sud, J. A. I. Pancher s.n.
evergreen trees. Branching sparse in remote pseudo- (holotype P). Fig. 21 133
whorls (Massarts model). Terminal buds absent.
Leaves spirally arranged, of two kinds: small and Podocarpus pectinatus Pancher ex Brongn. & Gris,
scale-like on leading and fertile shoots, larger Bull. Soc. Bot. France 16: 330. 1869.
and foliate on lateral, vegetative shoots; the larger Acmopyle alba J. T. Buchholz, Bull. Mus. Hist. Nat.
leaves bilaterally flattened, falcate-linear, spreading (Paris), sr. 2, 21: 281. 1949.
obliquely or in pinnate ranks on shaded shoots, sin-
gle-veined. Pollen cones lateral or terminal, axillary, Etymology
solitary or a few together, catkin-like; microsporo-
phylls helically attached to a slender rachis on very This species was named after J. A. I. Pancher, who
short stalks, triangular, with two basal pollen sacs collected the type specimen in 1869.
containing bisaccate pollen. Seed cones solitary or
occasionally grouped, on scale-leaved pedunculate Vernacular names
branchlets axillary or sub-terminal on leafy shoots,
when mature forming an irregular, fleshy and ver- No common names are recorded for this species.
rucose receptacle from several unfertilized bracts
which remain partly visible. Seeds single, from a Description
subterminal, inverted ovule, becoming (nearly) erect
at maturity, entirely enclosed by a fleshy epimatium, Small to medium size trees to 25 m tall, usually
its base partly enclosed by the receptacle, bluish pru- smaller; trunk monopodial, to 50cm d.b.h. Bark on
inose when mature. trunk becoming hard and scaly, breaking into small
plates, brown weathering grey; inner bark more
2 species or less fibrous. Branches spreading horizontally in
young trees, irregular and assurgent in older trees,
Distribution with foliage towards the ends. Foliage branches terete,
with small spreading leaf apices terminating spirally
New Caledonia; Fiji (Viti Levu). arranged, decurrent scale leaves; ultimate foliage
branchlets 315cm long, alternating, mostly plagio-
Key to the species of Acmopyle tropic, becoming deciduous after a few years, with
short scale leaves at the base which gradually enlarge
Leaves falcate to weakly S-curved, in middle to pectinately arranged foliage leaves. Foliage leaves
of branchlets 1030mm long, 1.53mm wide. on seedlings alternate, pectinate, linear, straight, in
Pollen cones elongating to 2025mm long, ca. the middle of the branchlet ca. 15mm long, 1.5mm
3mm wide. Fully developed receptacles 1520 wide, acute or pungent. Foliage leaves on young
810mm. New Caledonia A. pancheri trees and mature trees similar, alternate, pectinate
Leaves straight or falcate to weakly S-curved, in at 6090, straight or more often falcate towards
middle of branchlets 1025mm long, (0.6)2 apex, or on vigorous branchlets weakly S-curved, in
4(4.8) mm wide. Pollen cones elongating to the middle of branchlets 1030mm long, 1.53mm
58mm long, ca. 1.5mm wide. Fully developed wide, gradually shorter especially towards the proxi-
receptacles 79 78mm. Fiji A. sahniana mal and less so towards the distal end of branchlets,
twisted and decurrent at base; margins revolute; Uses
apex obtuse or acute, curved forward; midrib nar-
row above, often faint, prominent below; leaf colour No uses have been recorded of this species and it is
lustrous dark green above, with two whitish bands in cultivation only in a few botanic gardens.
separated by a green midrib and with green mar-
gins below. Stomata on both surfaces, in numerous
intermittent lines, some on the midrib on the lower Acmopyle sahniana J. T. Buchholz & N. E. Gray,
(abaxial) side and a few stomata near the base and/ J. Arnold Arbor. 28: 142. 1947. Type: Fiji: Western
or apex on the upperside. Pollen cones subtermi- Division, Viti Levu, Vakarogasiu Mountain,
nal or terminal on lateral foliage shoots, solitary [Namosi Province], J. W. Gillespie 3273
134 or 23 together, on short, scale-leaved peduncles, (holotype A).
initially globose, elongating to 2025mm long, ca.
3mm wide at anthesis; microsporophylls imbricate, Etymology
carinate, acute at first then acuminate, speading but
apically incurved, with two basal, small pollen sacs. This species was named after Prof. Birbal Sahni from
Seed cones subterminal or terminal on lateral foliage Lucknow, India, who studied the morphology of the
shoots, sometimes axillary to scale leaves or foliage genus Acmopyle.
leaves, solitary or sometimes 23 together, on up to
12mm long, curved, scale-leaved peduncles; com- Vernacular names
posed of several sterile and 12 fertile, amalgamated
bracts, forming a fleshy, swollen, irregularly shaped, No common names have been recorded for this
verrucose and pruinose receptacle 1520mm long species.
and 810mm wide. Seeds 12 at the distal end of the
receptacle, subglobose, 710mm diam. including Description
the covering epimatium, crested and pruinose when
full-grown. Small (monoecious?) trees to 12 m tall; trunk nor-
mally monopodial, to 20cm d.b.h. Bark mostly
Distribution smooth, sometimes pustulate, brown weather-
ing grey; inner bark red and more or less fibrous.
New Caledonia (Grande Terre). Branches sparse, spreading to form a crown half as
TDWG codes: 60 NWC wide as the tree height, with foliage towards the ends.
Foliage branches terete, with very small spreading
Ecology leaf apices terminating spirally arranged, decurrent
scale leaves; ultimate foliage branchlets 26(12)
Acmopyle pancheri occurs scattered in rainforest as cm long, alternating, mostly plagiotropic, becoming
an understorey tree and in low forest and vegeta- deciduous after a few years, with short scale leaves
tion bordering on maquis minier as a small canopy at base which often abruptly change to pectinately
tree. Its altitudinal range is from near sea level to at arranged foliage leaves. Foliage leaves on young
least 1200 m a.s.l. and it is found on ultramafic soil trees and mature trees similar, alternate, pectinate at
derived from serpentine or similar rock as well as 6090, straight or more often falcate towards apex,
on acidic soil from metamorphic schist. It is asso- or on vigorous branchlets weakly S-curved, in the
ciated with other conifers such as Araucaria spp., middle of branchlets 1025mm long, (0.6)24(
Dacrydium araucarioides and Agathis ovata, as well 4.8) mm wide, smallest towards proximal and dis-
as with numerous angiosperms. tal end of branchlets, twisted and decurrent at base;
margins revolute; apex acute, curved forward; mid-
Conservation rib narrow above, often faint, prominent below; leaf
colour lustrous dark green above, with two whitish
IUCN: NT bands separated by a green midrib and with green
margins below. Stomata on both surfaces, in numer- Ecology
ous intermittent lines, some on the midrib on the
lower (abaxial) side and a few stomata near the base Acmopyle sahniana is a small tree occurring sparsely
and/or apex on the upperside. Pollen cones subter- in low rainforest on mountain ridges and summits
minal or terminal on lateral foliage shoots, solitary in three localities on the island of Viti Levu. The alti-
or in pairs, sessile or on very short peduncles, ini- tudinal range is from 600 m to 1050 m a.s.l.
tially globose, elongating to 58mm long, ca. 1.5mm
wide at anthesis; microsporophylls imbricate, cari- Conservation
nate, acute, speading, with two basal, small pollen
sacs. Seed cones subterminal or terminal on lateral The very limited area of occupancy (AOO) of this
foliage shoots, sometimes axillary to scale leaves or species and an estimated population not exceed- 135
foliage leaves, solitary, on up to 6mm long, curved, ing 50 mature trees put this species in the Critically
scale-leaved peduncles; composed of 23 sterile and Endangered category. Proposed mining activities in
1(2) fertile amalgamated bracts, forming a fleshy, the area, which is not protected, further threaten this
swollen, irregularly shaped, verrucose and green or species (Doyle in Farjon & Page, 1999).
purple receptacle 79mm long and 78mm wide. IUCN: CR [C2a (i)]
Seeds solitary at the distal end of the receptacle,
ovoid, 79mm long, 56mm wide including the Uses
covering epimatium, striated and greyish violet with
whitish bloom when full-grown. No uses have been recorded of this species.
Distribution
Greek: actino- = rayed, star-like; strobus = cone; 2a. Seed cone scales with straight, acute apex
referring to the six scales coming together at the A. arenarius
apex of the seed cone. 2b. Seed cone scales with incurved, obtuse apex
A. pyramidalis
Description
136 Evergreen, monoecious (decumbent) shrubs or Actinostrobus acuminatus Parl., Index Sem. Hort.
trees with smooth, thin, flaking bark. Resin cavi- Florent. 1862: 25. 1862. Type: Australia: Western
ties in leaves. Branches short, stiff, spreading or Australia, [between Moore & Murchison Rivers;
ascending, forming a conical, pyramidal or bushy Swan River], J. Drummond 225 (holotype FI).
crown. Fastigiate forms common in nature. Leaves
in whorls of 3, decurrent; juvenile acicular leaves on Etymology
young plants only or also on mature plants; adult
leaves mostly shorter than 5mm, linear-lanceolate, The species epithet refers to the shape of the cone
abaxially keeled, denticulate on margins and keel apex (Latin: acuminatus = tapering from inwardly
or only on margins, acuminate-pungent, green or curved sides to a narrow point).
glaucous green, amphistomatic, stomata in 2 lines
on each face. Pollen cones small, cylindrical; micro- Vernacular names
sporophylls 1018, in whorls of 3, with 24 abaxial
pollen sacs. Seed cones solitary, with 23 together or Dwarf Cypress
aggregated along branches and stems, when closed
broadly globose with dome-shaped apex, broadly Description
globose-conical, or broadly utriculate. Bract-scale
complexes in two whorls of 3, of nearly equal size Shrubs, erect or decumbent to ascending, dense, 14.5
at maturity, subtended by 46 alternating whorls of m tall; trunk short or multistemmed. Bark smooth,
3 broad, imbricate scale leaves. Bract tips entirely soon flaking, thin, brown-grey. Branches numerous,
included in cone scales. Cone scales oblong, open- spreading or prostrate, persistent, forming a broadly
ing valvately, with light coloured seed scars towards conical or more irregular crown (often partly bur-
base. Columella a strong, acute spike. Ovules in 2 ied in blown sand). Foliage branches numerous,
whorls of 46 axillary to bracts, erect. Seeds 812 contorted, spreading irregularly or ascending, rigid,
per cone, with 3 wings. Seedlings with 2 cotyledons. slender, short, rather sturdy, angular with decur-
rent leaf bases, persistent, grey-brown to grey when
3 species leaves weather away. Leaves in alternating whorls of
3, decurrent; juvenile leaves on young and mature
Distribution plants, acicular, 1020 0.81.5mm, with spreading
free part (patent), abaxially keeled, acute-pungent,
SW Western Australia, in a narrow coastal strip light green or yellowish green, adult leaves linear-
from S of Shark Bay to near Albany. lanceolate, 24 1mm, with recurved or incurved
free apex, abaxially keeled, denticulate on margins,
Key to the species of Actinostrobus acuminate, green; stomata in 2 narrow lines on each
face. Pollen cones terminal on ultimate branchlets,
1a. Both juvenile, long acicular and adult, scale yellowish green turning light brown, 710 23mm;
leaves present on mature plants. Seed cones not microsporophylls 1016, in whorls of 3, ovate with
in clusters on stems A. acuminatus acute apex, slightly keeled abaxially, with 34 abaxial
1b. Leaves on mature plants short, scale-like. Seed pollen sacs. Seed cones terminal on short (ca. 1cm),
cones in clusters on stems 2 slightly thickened, lateral, short-leaved branchlets,
mostly solitary or with 23 together, when closed Actinostrobus arenarius C. A. Gardner, J. Roy.
broadly utriculate, with a distinct neck below apex Soc. W. Austral. 47: 54. 1964. Actinostrobus pyrami-
of scales, 1525 1020mm, glaucous green matur- dalis Miq. var. arenarius (C. A. Gardner) Silba,
ing to brown. Bract-scale complexes whorled, 6, Phytologia Mem. 7: 11. 1984; Actinostrobus pyrami-
of nearly equal size at maturity, subtended by 46 dalis Miq. subsp. arenarius (C. A. Gardner) Silba, J.
alternating whorls of 3 broad, imbricate scale leaves Int. Conifer Preserv. Soc. 13 (1): 1. 2006.
with denticulate margins; apex free, acuminate; Type: Australia: Western Australia, Tammin,
smaller lower whorls partly overlapping larger upper C. A. Gardner 610 (holotype PERTH). Fig. 22, 23
whorls. Bract tips entirely included in cone scales.
Cone scales oblong, 1520 510mm, more or less Etymology
concave and finely wrinkled, dark brown abaxially, 137
with recurved, acute apex, with smooth adaxial face, The species epithet refers to its habitat (Latin: arena =
blackish or purplish black, with light coloured seed sand).
scars towards base. Columella a strong, acute central
spike. Seeds 812 per cone, only a small number fer- Vernacular names
tile, about half of these smaller and situated between
bases of cone scales, irregularly triangular/angular, Bruce cypress
56mm long (including wings 912mm), yellowish
brown; wings narrow, 23mm wide. Description
Latin: afro- = African; carpus = fruit; name given met with universal recognition, in particular among
to distinguish the genus from the related genus South African botanists. Some morphological dis-
Podocarpus. tinctions stressed by Page, such as the lack of a succu-
lent and colourful receptacle, present in Podocarpus
140 Description sensu stricto, are in reality less rigidly distinct, with
the situation found in P. henkelii in particular rep-
Evergreen, dioecious, large trees. Resin canals (1) in resenting an intermediate stage. The function of the
leaves only. Bark thin, becoming scaly with small receptacle, serving seed dispersal by means of offer-
plates. Terminal buds small or absent. Leaves spirally ing birds an attractive imitation fruit, is apparently
inserted or (on young plants) opposite, twisted in taken over by the fleshy epimatium surrounding the
opposite directions so as to orientate the blades [on seed in Afrocarpus. There are botanists who would
opposite sides of the same shoot] with the adaxial side be inclined to emphasize such adaptational traits as
uppermost and the adaxial side downwards, pecti- significant, but others who would argue that they
nately arranged or ascending, flattened, coriaceous, do not necessarily indicate phylogeny (adaptation
narrowly lanceolate-elliptic to linear-lanceolate, with a can lead to convergent evolution) and if not, should
single midrib. Stomata on both surfaces (leaves amphi- be disregarded. Other morphological differences,
stomatic). Pollen cones axillary, solitary or in groups such as the amphistomatic leaves of the species in
of 23 on short naked peduncles, becoming narrowly Afrocarpus, appear to be more consistent and have
cylindrical and catkin-like; microsporophylls spirally supported a phylogeny separating Afrocarpus (Kelch,
inserted, with two pollen sacs containing bisaccate 1997). Recently, research into this taxonomic ques-
pollen. Seed cones axillary or just below foliage leaves tion has involved molecular (DNA) analysis (Barker
on a short peduncle with or without small scale leaves, et al., 2004 and papers cited therein) and these data
consisting of a few sterile bracts and one larger fer- also gave support to the recognition of Afrocarpus
tile bract with an axillary, inverted ovule. Seeds single as distinct from Podocarpus. Cladistic analyses of
per reduced cone, subtended by small, withering morphological as well as molecular data therefore
scales (not by a receptacle), entirely enclosed by a appear to confirm the taxonomy proposed by Page,
fleshy, subglobose to obovoid or ellipsoid epimatium, as is here adopted.
maturing from greenish to yellow or reddish brown;
seed proper with a hard, strongly sclerified seed coat.
Key to the species of Afrocarpus
5 species
The species of Afrocarpus are difficult to key out on
Distribution vegetative characters only; it is therefore necessary
to examine especially the seeds with and without
E Africa: Ethiopia, Kenya, Uganda, Rwanda, their soft covering (epimatium) and in their fully
Burundi, Congo Republic (Kivu), Tanzania, Malawi; mature state in most cases.
S Africa: South Africa, Mozambique (Loureno
Marques), Swaziland; W Africa: So Thom. 1a. Adult leaves 37(8) mm wide, up to 11 cm long
(usually to 8 cm long). Microsporophylls of
Taxonomic notes pollen cones broadly triangular A. mannii
1b. Adult leaves (1.5)24(5) mm wide, up to 8 cm
The genus Afrocarpus, raised to that rank by Page long (usually to 5 cm long). Microsporophylls
(1989) from a section under Podocarpus, has not of pollen cones triangular-trullate 2
2a. Buds very small, 0.61 mm diam. Midrib rounded with or without a rostrate apex. Leaves on
prominent on both sides of leaves. Seed coat seedlings and young plants mostly opposite, nar-
(3)46(8) mm thick (remove epimatium) rowly linear-lanceolate, up to 17 cm long and 48
A. usambarensis mm wide, straight or falcate, tapering to a fine point.
2b. Buds larger, ca. 2 mm diam. Midrib prominent Adult leaves shorter, (2)35(6) cm long, (1.5)2
on the adaxial (upper) sides of leaves only. Seed 4(5) mm wide, spirally arranged, twisted at the
coat 14 mm tick (remove epimatium) 3 attenuate base, spreading to ascending, straight or
3a. Seeds including the epimatium 2030 mm rarely slightly falcate, linear-elliptic or with parallel
long; seed proper (remove epimatium) rugose, sides, gradually or abruptly tapered above 3/4 of their
with a 24 mm thick seed coat A. dawei length, with a raised midrib most prominently on the
3b. Seeds including the epimatium 1220(23) mm adaxial (lower) side, grey-green; apex acute. Stomata 141
long; seed proper smooth or verrucose, with a on both surfaces, arranged in numerous intermittent
11.5 mm thick seed coat 4 lines not well separated by the midrib. Pollen cones
4a. Adult leaves (1)24(4.5) cm long. Seed proper solitary or with 23 on very short stalks or subses-
spherical/compressed, 1014 mm diam., with a sile, axillary to foliage leaves or not, subtended by a
verrucose seed coat A. falcatus whorl of papery bracts, initially subglobose, elongat-
4b. Adult leaves (1.5)36(8) cm long. Seed proper ing to cylindrical, 1020(25) mm long, 2.53.5 mm
ovoid/compressed, 1218 mm long, with a diam.; microsporophylls spirally arranged, imbricate
smooth seed coat A. gracilior before anthesis, triangular-trullate, ca. 1 mm wide,
with denticulate-lacerate margins and acute or apic-
ulate apex, bearing two subglobose pollen sacs. Seed
Afrocarpus dawei (Stapf) C. N. Page, Notes Roy. cones solitary on small, scaly or leafy branchlets situ-
Bot. Gard. Edinburgh 45: 384. 1989. Podocarpus ated axillary to or below foliage leaves, with several
dawei Stapf, in Prain, Fl. Trop. Afrika 6 (2): 342. sterile and one terminal, larger fertile bract. Mature
1917. Type: Uganda: Nile Land, South Buddu, seed cones with a single seed subtended by a single,
Kaigera River, M. T. Dawe 961 (holotype K) short bract; with seed enclosed by a fleshy, firm epi-
[holotype K, not found; isotype B, destroyed]. matium that ripens from green or glaucous green to
yellow, globose or subglobose, 2530 mm long. Seed
Etymology proper ovoid but slightly compressed laterally, 1621
mm long, 1215 mm wide, with a rugose surface and
The species epithet commemorates the botanist hard, 24 mm thick seed coat.
Morley Thomas Dawe (18801943), who collected
the type specimen. Distribution
This species does not fulfil the criteria for V, E, Bastard yellowwood, Outeniqua yellowwood;
CR based on GIS information; its extent of occur- Outeniekwageelhout; inkoba (South Africa);
rence (EOO) is 175,410 km (possible NT) and area umgeya (Xhosa, Zulu)
of occupancy (AOO) is 3,200 km (NT). Human
Footprint AVG in the area is low (7). It is considered Description
NT on the basis of observational reports on exploita-
tion cited on herbarium specimens, which indicates Trees generally to 25 m tall but attaining 60 m,
there is reason to suspect decline under criterion A2, with a massive trunk, d.b.h. to 2 m. Bark smooth in
142 but it cannot be quantified. A notable population young trees, flaking in rectangular or rounded small
occurs in the Minziro Forest Reserve in Tanzania, plates in large trees, purplish brown or dark brown,
where 16 of 32 (=50%) herbarium collections used in weathering grey. Branches ascending and spreading
the assessment were made. forming a broad, domed crown. Foliage dense, on
IUCN: NT numerous branches; new lateral branchlets ridged or
more or less quadrangular; terminal buds small, ca.
Uses 2 1 mm, or absent; bud scales narrowly triangu-
lar, acute. Leaves spirally arranged, on seedlings and
The timber of this species, which can grow a tall, young plants narrowly linear-lanceolate, up to 12 cm
straight bole without branches, is valued for con- long and 36 mm wide, straight or falcate, tapering
struction and carpentry or joinery work, and trees to a fine point. Adult leaves much shorter, (1)24(
are singled out despite the seasonally difficult access 4.5) cm long, (1.2)24(5) mm wide, twisted at the
to be logged. It is not exported and used locally narrowed base, spreading to ascending, straight or
or regionally. The species is not known to be in slightly falcate, linear-lanceolate to linear-elliptic,
cultivation. with a conspicuously raised midrib adaxially (lower
side), obscurely present abaxially, grey-green; apex
Afrocarpus falcatus (Thunb.) C. N. Page, Notes Roy. acute to obtuse. Stomata on both surfaces, arranged
Bot. Gard. Edinburgh 45: 383. 1989. Taxus falcata in numerous intermittent lines not well separated
Thunb., Prodr. Pl. Cap.: 117. 1800; Podocarpus fal- by the midrib. Pollen cones solitary or with 24
catus (Thunb.) Endl., Syn. Conif.: 219. 1847; Nageia on very short stalks or subsessile, axillary to foli-
falcata (Thunb.) Kuntze, Revis. Gen. Pl. 2: 800. age leaves or not, subtended by a whorl of papery
1891 (nom. illeg. Art. 53.1); Decussocarpus falcatus bracts, initially subglobose, elongating to cylindri-
(Thunb.) de Laub., J. Arnold Arbor. 50: 359. 1969. cal, 513(15) mm long, 23 mm diam.; microspo-
Type: South Africa: Cape Province, [e Cap. b. Sp. rophylls spirally arranged, imbricate before anthesis,
Ribeek(capel?), Vleermuysdrift], C. P. Thunberg broadly triangular-trullate, ca. 0.6 1 mm, with lac-
UPS 23779 (holotype UPS). Fig. 24 erate margins and acute or apiculate apex, bearing
two slightly elongate pollen sacs. Seed cones soli-
Podocarpus gracilimus Stapf, in Prain, Fl. Trop. tary on small, scaly branchlets situated axillary to
Africa 6 (2): 343. 1917. or below foliage leaves, with several sterile and one
terminal, larger fertile bract. Mature seed cones with
Afrocarpus gaussenii (Woltz) C. N. Page, Notes Roy. a single seed subtended by a single, short bract; with
Bot. Gard. Edinburgh 45: 384. 1989; Podocarpus seed enclosed by a fleshy, firm epimatium that rip-
gaussenii Woltz, Trav. Lab. Forest. Toulouse T. 1 (8, 2): ens from glaucous green to yellow or light reddish
6. 1969. brown, globose to obovoid, 1218 mm long, resin-
ous. Seed proper nearly spherical but slightly com-
Etymology pressed laterally, 1014 mm diam. with a verrucose
surface and a 1 mm thick, hard seed coat.
The species epithet refers to the (occasionally) sickle-
shaped leaves.
Taxonomic notes uals or occasionally form a small group. Presumably
their regeneration is dependent on episodal canopy
In southern Africa, this species is still classified disturbance similar to e.g. Podocarpus totara in New
under Podocarpus (see e.g. Keith Coates Palgraves Zealand and many other big, long-lived conifers.
popular handbook Trees of Southern Africa in its
several editions, 19771988) despite obvious differ- Conservation
ences in the lack of development of a receptacle in
the seed cone and in the anatomy and morphology of IUCN: LC
the leaves, e.g. amphistomatic versus hypostomatic
leaves. Other differences are the placement of pollen Uses
cones and seed cones below foliage leaves on scaly 143
dwarf shoots and the thickening and colouring of The wood of Afrocarpus falcatus, incorrectly known
the epimatium to resemble a yellow plum-like fruit. as yellowwood (= Podocarpus latifolius), is valuable
None of these characters are shared with Podocarpus especially in the large sizes it attains in the south-
sensu stricto anywhere in the world. A form with ern Cape. It was used in the past for ship masts and
small, very narrow leaves (1.52.5 cm 1.52 mm) is still in high demand for boat building. The sawn
from Transvaal, South Africa was described as a new timber is also used in construction for beams and
species Podocarpus gracillimus by Stapf, but there is rafters, house floors and wall panelling, carpentry
much variation in leaf sizes, which may in part be and joinery, and furniture making. In horticulture
influenced by growing conditions. The taxon origi- it is increasingly popular as an amenity tree, mainly
nally described as P. gaussenii from Madagascar has in countries with a mild climate like South Africa,
turned out to be a case of introduction of A. falcatus e.g. western USA, Australia and New Zealand. It can
to that island; it has not been found growing in the also be seen in several botanic gardens and arboreta
wild in recent surveys of the forest flora. outside South Africa.
Distribution
Afrocarpus gracilior (Pilg.) C. N. Page, Notes Roy.
SE & S Africa: from Malawi and Mozambique to Bot. Gard. Edinburgh 45: 383. 1989. Podocarpus
Kwazulu Natal and Eastern and Western Cape gracilior Pilg., in Engler, Pflanzenr. IV.5 [18]: 71.
Provinces, South Africa. 1903; Decussocarpus gracilior (Pilg.) de Laub., J.
TDWG codes: 26 MLW MOZ 27 CPP-EC CPP-WC Arnold Arbor. 50: 359. 1969. Type: Ethiopia: [Gerra
NAT OFS SWZ TVL-GA TVL-MP TVL-NP Abuna Tekla Zlaimanot (Haimanot), auf Bergen
2500 m . M.], G. H. W. Schimper 1160 (syntype K).
Ecology Fig. 25, 26
pinheiro de So Thom, pinheiro da terra Afrocarpus mannii is endemic on the volcano Pico
(Portuguese) de So Tom from ca. 1450 m to the summit area at
2142 m a.s.l. It is nowhere a tall tree and at the sum-
Description mit it is reduced to dwarfed krummholz. It is com-
mon in the high montane cloud forest where this has
Trees generally to 15 m tall but on the summit area as remained undisturbed.
krummholz. Bark undescribed. Branches ascending
and spreading forming a broad crown. Foliage rela- Conservation
tively sparse; new lateral branchlets ridged or more
or less quadrangular; terminal buds small, ca. 2 Deforestation at lower to middle altitudes on the
1 mm, or absent; bud scales triangular, acuminate. mountain is the main threat to this species, which is
Leaves spirally arranged, on seedlings and young endemic to the island.
plants linear-lanceolate to subfalcate, up to 16 cm IUCN: VU (D2)
long and 48 mm wide, straight or falcate, tapering
to a fine point. Adult leaves slightly shorter, (2)3 Uses
8(11) cm long, 37(8) mm wide, twisted at the
petiolate base, spreading, straight or slightly falcate, The timber of Podocarpus mannii is valuable in trees
lanceolate to linear-lanceolate, with a conspicuously of good size and shape, which have become scarce. It
raised midrib adaxially (lower side) and obscurely is used for light construction. This species has been
present abaxially, grey-green; apex acute to obtuse. planted in rural areas in Cameroon and Ivory Coast
Stomata on both surfaces, arranged in numerous and probably elsewhere in W Africa as a canopy tree
intermittent lines not well separated by the mid- or windbreak for coffee plantations and as an ame-
rib. Pollen cones solitary or with 2, sessile, axillary nity tree in villages.
Afrocarpus usambarensis (Pilg.) C. N. Page, lets situated axillary to or below foliage leaves, with
Notes Roy. Bot. Gard. Edinburgh 45: 384. 1989. several sterile and one terminal, larger fertile bract.
Podocarpus usambarensis Pilg., in Engler, Pflanzenr. Mature seed cones with a single seed subtended
IV.5 [18]: 70. 1903. Type: Tanzania: Tanga Prov., by a single, short bract; seed enclosed by a fleshy,
Usambara Mts., Mtai Hill, near Mpare village, firm epimatium that ripens from green or glaucous
C. Holst 2467 (syntype K). green to yellow, globose or sometimes broad ellip-
soid, (17)2330(35) mm long. Seed proper globose
Etymology or broadly ellipsoid but slightly compressed later-
ally, (15)2025(30) mm long, with a rugose-pus-
The species epithet refers to the Usambara Mountains ticulate surface and a (3)46(8) mm thick, hard
146 from where it was first described. seed coat.
mse, muze (Usambara Mts., Tanzania) Tanzania, Kenya (Kyulu Hills, Taita Taveta District).
TDWG codes: 25 KEN TAN
Description
Ecology
Trees to 30 m tall, d.b.h. to 2 m. Bark smooth in
young trees, flaking in rectangular or rounded small Afrocarpus usambarensis occurs in montane ever-
plates in large trees, dark brown, weathering grey. green rainforest and dry evergreen forest, mixed
Branches ascending and spreading forming a broad, with co-dominant angiosperms. Elevation ranges
domed crown. Foliage dense, on numerous branches; from ca. 1500 m to ca. 3000 m a.s.l. Trees are often
new lateral branchlets ridged or more or less quad- solitary but not emergent, only reaching into the
rangular; terminal buds very small, 0.61 mm diam., general canopy of the forest. In rainforest it occurs
or absent; bud scales rounded with or without an often with Podocarpus milanjianus; the co-dom-
apiculate apex. Leaves on seedlings and young plants inant angiosperm tree in this wetter forest type is
mostly opposite, narrowly linear-lanceolate, up to 13 often Ocotea usambarensis, but many other species
cm long and 47 mm wide, straight or falcate, taper- may occur with it. In dryer evergreen forest Olea and
ing to a fine point. Adult leaves shorter, (1.5)35(6) Ficus are common associates of A. usambarensis,
cm long, (1.5)24(5) mm wide, spirally arranged, other taxa are e.g. Calodendrum capense, Syzygium
twisted at the narrowed base, spreading to ascend- cordatum and Bridelia micrantha. These drier forests
ing, straight or rarely slightly falcate, linear-elliptic are often degraded or converted to coarse grassland
or with parallel sides, gradually tapered above 2/3 or in which A. usambarensis can survive as isolated
3/4 of their length, with a raised midrib usually pres- trees, at least for a time.
ent on both sides, grey-green; apex acute. Stomata
on both surfaces, arranged in numerous intermittent Conservation
lines not well separated by the midrib. Pollen cones
solitary or with 23 on very short stalks or subsessile, This species is considered threatened based on direct
axillary to foliage leaves or not, subtended by a whorl observation of intense local exploitation (aerial pho-
of papery bracts, initially subglobose, elongating to tography of many saw pits both within and with-
cylindrical, (5)1020(26) mm long, 2.53.5 mm out reserves). This species is under severe threat
diam.; microsporophylls spirally arranged, imbri- from illegal logging in the Chome Forest Reserve
cate before anthesis, triangular-trullate, ca. 0.8 mm in Tanzania (evidence from aerial photography);
wide, with denticulate-lacerate margins and acute or the same type of saw pit exploitation is known from
apiculate apex, bearing two subglobose pollen sacs. other locations. General deforestation and fires
Seed cones solitary on small, scaly or leafy branch- are also reducing the rainforest, which is usually
limited in extend even naturally. This species is the Uses
most valuable and specifically targeted tree for (ille-
gal) logging in this type of forest. It is present in the This species, yielding yellowwood or podocarp
following reserves in Tanzania: Chome, Hanang, wood is highly valued for its timber and exploited
Mafwomero, Mkusu, Nou, Shagayu and Wotta. The mainly for sawn timber used in construction of
evidence is that this does not prevent illegal logging houses. The wood is yellowish in colour, straight-
on a large scale using sawpits to remove and process grained, and clean of knots and can be used for gen-
individual trees. eral carpentry and furniture as well. This species is
IUCN: EN [A2, A4 + B 2ab (ii, iii, iv, v)] not known to be in cultivation.
147
Agathis Salisb., Trans. Linn. Soc. London 8: 311. 1807 (nom. cons.). Dammara
(Rumph.) Lam., Enum. Pl. Hort. Berol. 2: 411. 1822. Type: Agathis dammara
(Lamb.) Rich. & A. Rich. [Dammara loranthifolia Link (Pinus dammara Lamb.)]
(Araucariaceae).
Salisburyodendron A. V. Bobrov & Melikyan, axis, the other opposite and rudimentary to some-
Komarovia 4: 62. 2006. Type: Salisburyodendron times virtually absent. Seedlings with 2 cotyledons.
australis (D. Don) A. V. Bobrov & Melikyan [Agathis
australis (D. Don) Lindl.]. 17 species
148
Greek Agathis = a clew or ball of thread; it refers to Distribution
the seed cone.
Malesia: Malay Peninsula, Sumatera, Borneo,
Description Sulawesi, Philippines, Maluku [Moluccas], New
Guinea, New Britain; Australia: coastal Queensland;
Evergreen monoecious trees, often of great size, SW Pacific: New Caledonia, Vanuatu, Fiji, Solomon
monopodial with straight boles. Resin canals in Islands (Santa Cruz Group), New Zealand (North
bark, leaves and seed cones. Branching in sub-ver- Island).
ticillate pseudo-whorls (Massarts or Rauhs model),
truncated in very large trees. Apical buds globose Taxonomic notes
with imbricate scales. Leaves subopposite to oppo-
site, short petiolate, broad and multi-veined, coria- Few attempts have been made to investigate the
ceous, more or less hypostomatic, extremely variable possible relationships among the various species
in shape and size within a single tree; those on young in this genus. Most studies of this kind have con-
trees usually larger than on mature trees. Pollen cones cerned themselves with relationships at the level
appearing after seed cones, axillary, solitary, sessile of genus. Phylogenetic relationships of taxa within
to pedunculate, subtended by more or less decussate the Araucariaceae were investigated by Gilmore
bracts, the lower pair of which may be leaf-like or & Hill (1997), Setoguchi et al. (1998) and Kershaw
not, elongating to a cylindrical or catkin-like shape & Wagstaff (2001) (the first two papers used rbcL
after anthesis. Microsporophylls helically attached to DNA markers) and none of these studies sampled
a rachis in imbricate or tesselate arrangement, con- all species. We therefore still lack a hypothesis of the
sisting of a short stalk and a more or less peltate head phylogeny of the species of Agathis upon which a
of varied shape according to species; bearing from classification of the genus could be based and, indeed,
212 elongated pollen sacs directed inward towards no infrageneric classification has been proposed. In
cone rachis. Seed cones axillary or sometimes termi- order to conveniently and more reliably key out the
nal, solitary on stout stalks, globose to obovoid, usu- species, they have here been divided into two geo-
ally smooth but sometimes rough with bossed scale graphically determined groups; each of these covers
margins, green or slightly glaucous, disintegrating roughly half of the entire range of the genus. No tax-
when mature and drying. Cone scales composed of onomy is implied with this division and it only serves
a fused bract and seed scale, the visible and largest the keys. The South-West Pacific here includes New
part made up of the bractaceous element, helically Zealand and Malesia is the region, as defined in Flora
attached to a stout rachis shorter than the cone, Malesiana, from Peninsular Malaysia to New Guinea
imbricate, thin except the outer, mostly exposed and the Solomon Islands. Few species are known in
margin. Seeds 1 per cone scale, inverted, more or less cultivation outside their own region as here defined
ovoid but strongly flattened, with two thin, membra- (the exceptions almost all involve A. australis from
nous wings, one obliquely placed relative to the seed New Zealand) and a key to cultivated species is for
this reason not considered necessary.
Key to the species of Agathis in Australia and 6b. Pollen cones with 616 decussate, imbricate or
the SW Pacific free, linear to triangular bract scales. Juvenile
leaves 1120 cm long; adult leaves (2.5)49 cm
Pollen cones provide most of the diagnostic charac- long 7
ters of species in this genus; under mature trees (all 7a. Pollen cones short cylindrical to nearly oval,
are monoecious) these are often found in the leaf litter 22.5 cm long, 810 mm wide; intact cones with
under their canopy. Only fully expanded cones should 2 leaf-like bracts subtending 68 free bract
be taken into account. Leaves are highly variable but scales A. lanceolata
differ between juvenile (phase 1) leaves and adult 7b. Pollen cones cylindrical, (2)2.55(7) cm long,
(phase 2) leaves as treated in the species descriptions; 615 mm wide; lacking 2 leaf-like bracts, with
only maximum length for the first category appears to 816 imbricate bract scales 8 149
be informative in this context. A single twig with foli- 8a. Bark on trunk with numerous small, granular
age can never be determined with the macroscopic lenticels. Bract scales at base of pollen cone 12
leaf characters (measurements and shapes) alone. mm wide; microsporophyll heads umbonate.
Seed cones up to 7 cm diam A. montana
1a. Pollen cones short cylindrical, 0.91.6 cm long, 8b. Bark on trunk without or with few lenticels.
48 mm wide 2 Bract scales at base of pollen cone 47 mm
1b. Pollen cones cylindrical or oval-fusiform, wide; microsporophyll heads nearly flat or con-
26(7) cm long, (6)718 mm wide 3 vex. Seed cones 913 cm diam 9
2a. Microsporophylls of pollen cones imbricate, 9a. Pollen cones with 816 decussate bract scales at
more or less convex. Seed cones 3.55 cm diam.; base, 69 mm wide; microsporophyll heads
cone scales with a bossed upper margin nearly flat with angular upper margin
A. atropurpurea A. moorei
2b. Microsporophylls of pollen cones tessellate, 9b. Pollen cones with up to 8 decussate bracts
prismatic. Seed cones 6.510 cm diam.; cone scales at base, 815 mm wide; microsporophyll
scales with a smooth, slightly rounded upper heads convex, often notched at apex
margin A. microstachya A. macrophylla
3a. Juvenile leaves to 6 cm long. Pollen cones on
(4)520 mm long, stout peduncles; basal bract Key to the species of Agathis in Malesia
scales short, rounded or triangular. 5
3b. Juvenile leaves to 20 cm long; pollen cones ses- 1a. Juvenile (phase 1) leaves to 8 cm long, adult
sile or on short peduncles to 7 mm long; basal (phase 2) leaves to 5 cm (occasionally to 67
bract scales 17 mm wide, linear to triangular cm) long, the shortest leaves often obovate or
4 oval-orbiculate (length less than 2 width) 2
4a. Bark exfoliating with small flakes (smaller than 1b. Juvenile (phase1) leaves to 14 cm long, adult
10 cm). Pollen cones 813 mm wide, with 810 (phase 2) leaves to 12(14) cm long, the shortest
decussate, imbricate basal bract scales leaves variable in shape but not obovate or oval-
A. robusta orbiculate 4
4b. Bark exfoliating with large flakes (to 15 cm). 2a. Pollen cones when fully expanded 1.21.5 cm
Pollen cones 1518 mm wide, with 46 decus- long, 56 mm wide, lacking 2 leaf-like bracts
sate, free basal bract scales A. silbae subtending the cluster of smaller bract scales
5a. Bark on trunk smooth, with small or large A. orbicula
flakes. Trees becoming very large 6 2b. Pollen cones when fully expanded 2.54 cm
5b. Bark on trunk fissured, rough and scaly. Trees long, 810 mm wide; intact cones usually with 2
often stunted, up to 10 m tall (rarely taller to 25 leaf-like bracts up to 3 cm long, subtending a
m in forest) A. ovata cluster of smaller bract scales 3
6a. Pollen cones with 6 decussate, imbricate, short 3a. Two subtending bracts of pollen cones to 30
rounded bracts scales. Juvenile leaves to 6 cm 10 mm. Seed cone scales with a boss on
long; adult leaves 2.34(7) cm long the upper margin. Endemic of Peninsular
A. australis Malaysia A. flavescens
3b. Two subtending bracts of pollen cones to 20 4 Description
mm. Seed cone scales lacking a boss on the
upper margin. Endemic of Borneo Trees to 50 m tall and 2.5 m or more d.b.h. with
A. kinabaluensis clear cylindrical bole to 30 m and a rounded or open
4a. Pollen cones when fully expanded 89 cm long, crown with spreading to ascending branches. Bark
2535 mm wide; short basal bracts in 2 decus- smooth or sometimes scaly, exfoliating in irregular
sate pairs; microsporophyll heads 56 48 thin scales of up to 15 cm across; outer bark dark
mm (length width) A. borneensis brown or red-brown, with purple patches under
4b. Pollen cones when fully expanded (2.2)36 newly fallen scales; inner bark reddish, exuding
(7) cm long, 813 mm wide; short basal bracts white resin. Leaves subopposite, thin or thick, multi-
150 in 25 decussate pairs; microsporophyll heads nerved, coriaceous, glabrous, light green, sometimes
0.72 12.5 mm (length width) 5 glaucous on the underside especially leaves on new
5a. Microsporophylls in tessellate arrangement, shoots. Leaves on saplings and young trees and/or
their heads prismatic; short basal bracts of pol- in shade linear-lanceolate to elliptic, 58 cm long,
len cones in 45 decussate pairs 6 1.53 cm wide; leaves in crowns of mature trees lan-
5b. Microsporophylls in imbricate arrangement, ceolate to elliptic, 37 cm long, (0.5)12 cm wide,
their heads more or less convex; short basal short petiolate and with an obtuse apex. Pollen cones
bracts of pollen cones in 24 decussate pairs 7 axillary, solitary on a 23 mm long, stout peduncle,
6a. Pollen cones when fully expanded barrel- short cylindrical, when full grown 0.91.6 cm long,
shaped, 2.23 cm long, 1012 mm wide. Bark on 47.5 mm diam., with 810 decussate (in 45 pairs),
large trunks with large (to 10 cm) flakes imbricate, 12.5 mm wide bract scales at their base.
A. labillardierei Microsporophylls in imbricate arrangement; head
6b. Pollen cones when fully expanded cylindrical, with rounded or minutely mucronate upper mar-
46(7) cm long, 813 mm wide. Bark on large gin, 0.60.8 mm wide, 0.50.7 mm high in mature
trunks with small (to 5 cm) flakes A. robusta cones, bearing 25 pollen sacs. Seed cones solitary
7a. Short basal bracts of pollen cones in 24 decus- on thick peduncles, globose, 3.55.5 cm long, 3.55
sate pairs, free spreading; peduncle absent or to cm wide, more or less rough, green and resinous,
5 mm long. Seed cone scales with smooth, ripening brown. Cone scales with thick, slightly
slightly rounded margins A. dammara bossed, incurved upper margins, 1.62.3 cm long,
7b. Short basal bracts of pollen cones in 23 decus- 2.33 cm wide, broad triangular to reniform with
sate pairs, imbricate; peduncle 210 mm long. rounded corners and more or less flanged on either
Seed cone scales with bossed upper margin side. Seeds 79 45 mm, ovoid, flattened, with two
A. lenticula unequal wings; largest wing 1215 8 mm; smallest
wing a small acute triangular point 2 mm long oppo-
Agathis atropurpurea B. Hyland, Brunonia 1 (1): site largest wing.
109. 1978. Type: Australia: Queensland, Cook
District, Bellenden Ker, [N.P.R. 226], B. P. M. Distribution
Hyland 5776 (holotype BRI).
Australia: NE Queensland (Cook District).
Etymology TDWG codes: 50 QLD-QU
Description Ecology
Trees to 55 m tall, to 5 m d.b.h. or more in veteran Agathis australis is the dominant tree in mixed coni-
trees, self-pruning and with a long, clear, almost fer-angiosperm subtropical lowland primary forest
that once covered the northern peninsulas of North man-made disturbance events. This regeneration is
Island from near sea level to 375 (600?) m a.s.l., but possible, in ecologically suitable locations, within
of which only small remnants remain today. It is not some 80,000 ha of existing forest reserves. The
clear why it did not occur naturally further south Department of Conservation in New Zealand has
on the island, as planted trees grow well there now. plans to establish a Kauri National Park in order to
Perhaps the dynamics of the Kauri forest in its natu- give full protection to these forests. If IUCN Red
ral state played a part in this limitation. The species List criteria were strictly applied to this history of
is dependent for successful regeneration on episodal reduction of population size (A criterion), A. aus-
disturbance of the forest by fire or storm removing tralis would fulfil the criteria for a listing as (at least)
sizable swathes of forest cover. Even-aged stands Endangered (EN), even though the causes of decline
152 of large trees in groves and a few scattered giant have now ceased, because the reduction amounts
trees are a common pattern in the Waipoua Forest, to more than 70% over the last three generations
the largest of the remaining primeaval Kauri forest (of mature trees). Conservationists in New Zealand
remnants. The giants are more than 1000 years old object to such a rating because to them it would
and have survived one or more disturbances, act- appear to be a denial of their successful efforts to
ing as seed trees. In such a mature stand of Kauri curb the destruction and save this iconic tree from
there is little regeneration. Forest succession with- extinction. On the other hand one could observe
out disturbance eventually leads to dominance of that most of the historical reduction will be perma-
angiosperms. Early phases in disturbed areas are nent under human occupation and land use and that
dominated by Leptospermum scoparium and Kunzea while the reduction has ceased (and is even being
ericoides (Myrtaceae) into which A. australis invades reversed) no one can guarantee what the priorities
abundantly. The conifer Phyllocladus trichomanoides of future generations are going to be.
arrives next. Via stages with increased tree species IUCN: NT
diversity and the establishment of more shade toler-
ant trees, above which the rickers, i.e. Kauri trees Uses
with conical habit, rise, the forest restores itself.
A drastic thinning of Kauri trees then results in The kauri of New Zealand was once the most impor-
fewer trees with ever wider spreading crowns, ris- tant timber tree of these southern islands, but in the
ing above the canopy as emergents. Eventually there short space of roughly 50 years this natural resource
is another disturbance, but these cycles may have a had nearly been exhausted through one of the most
duration of 500 years or longer. Common conifers wanton campaigns of exploitation and short-term
in mature Kauri stands are Dacrydium cupressinum, thinking in the history of European colonisation
Prumnopitys ferruginea, P. taxifolia, Podocarpus of foreign lands. For several decades, the protec-
totara and Dacrycarpus dacrydioides; common co- tion of remaining forests and regeneration projects
dominant angiosperms are e.g. Beilschmiedia spp., have now ensured its continuing existence, but the
Weinmannia racemosa and Metrosideros umbellata. great expanse of kauri forest has nevertheless been
greatly reduced. The timber of old growth stands is/
Conservation was of large dimensions and the properties of the
wood are excellent for joiners work, boat building
Since the arrival of Europeans in New Zealand, the and carpentry. Black kauri, dark brown in colour
estimated 1,215,000 ha once covered by primeaval and very hard and durable, came from logs buried in
kauri forests have been reduced to ca. 7,500 ha. peat deposits. Wood from burls is beautifully figured
These remnants are now strictly protected; the larg- and was used in furniture and for panelling. The vast
est reserve is the Waipoua Forest covering 9,105 ha abundance of timber in the heyday of exploitation
(not all of it is dominated by A. australis) created in meant that it was also put to less refined use, such
1952. However, natural regeneration, where land as mine-props and railway sleepers. Another valu-
use as mixed (semi-)natural forest has not changed, able product is resin (copal) of which this tree pro-
is abundant in many places because the species of duces great quantities. During and after the timber
course does not distinguish between natural and exploitation, a veritable copal rush took place, with
thousands of resin diggers coming from all over to anthesis to 89 cm long and up to 35 mm wide, with
dig up the semi-fossil resin in the cut-over forests. 4 decussate (in 2 pairs) 58 mm wide, free spreading
Much of this was used in the manufacture of lino- bract scales at their base. Microsporophylls in imbri-
leum, paint and varnish. This species has been taken cate arrangement; head slightly convex towards the
into cultivation both in (modest scale) forestry plan- upper, rounded, paler coloured, minutely erose-den-
tations and as an ornamental tree in several coun- ticulate margin, 48 mm wide and 56 mm high in
tries with a mild climate. mature cones, bearing 410 oblong pollen sacs. Seed
cones solitary on thick peduncles, broadly ellipsoid
to globose, to 1013 cm diam., smooth, green, res-
Agathis borneensis Warb., Monsunia 1: 184. 1900. inous, ripening brown. Cone scales with slightly
Type: Malaysia: Sarawak, [locality not stated], O. rounded, incurved or slightly projecting upper mar- 153
Beccari 596 (syntype K). Pl. 5, Fig. 30 gins, ca. 3.5 cm long, 3.54.5 cm wide in larger cones,
roughly triangular with rounded corners and more
Agathis endertii Meijer Drees, Bull. Jard. Bot. or less flanged on either side. Seeds 1215 78 mm,
Buitenzorg, ser. 3, 16: 470. 1940. ovoid-oblong, with two unequal wings; largest wing
ca. 20 13 mm; smallest wing a small blunt triangle
Etymology 35 mm wide opposite largest wing.
plate 5. Agathis borneensis. 1. Habit of tree. 2. Foliage branch. 3. Leaf. 4. Pollen cone. 5. Microsporophylls.
6. Immature pollen cone. 7. Seed cone.
Ecology boats, and panelling to veneer and tool or furni-
ture making. Drawing boards are made of its wood
Agathis borneensis occurs in lowland to upland trop- as it is extremely easy to plain to a smooth surface.
ical rainforest as scattered emergent trees and in low Its odourless quality was noted in the manufacture
lying kerangas forest on sandy or sometimes peaty of food containers, until plastics took over from it.
soils, where it can form extensive pure stands, or The inner bark exudes a translucent to white resin
occurs mixed with the following conifers: Dacrydium known as copal and is still used for varnishes in
pectinatum, Falcatifolium falciforme, Nageia wallichi- photographic colour prints and as a component for
ana, Podocarpus spp., and Sundacarpus amarus. The the paint used to make lines etc. on tarmack road
most common angiosperm tree on peaty soils grow- surfaces. There is still a large export trade in its tim-
ing with A. borneensis is probably Gonystylus banca- ber, but with a deminishing quantity per annum and 155
nus (Thymaelaeaceae). In lowland to lower montane a trend to shift from round wood to sawn timber,
rainforest it can be associated with Dipterocarpaceae which fetches much higher prizes. This species (and
and/or Fagaceae; however, Agathis often retreats to A. dammara) are planted on a fairly large scale in for-
ridges with thin, rocky soils or to water-logged areas estry plantations in Jawa, but only locally on a small
where these dominant angiosperms are less vigor- scale within its native range. This will have to change
ous. It is also reported from heath forest which dramatically if the resource is to be made anywhere
occurs on higher mountain ridges and summits and near sustainable for the future. Agathis borneensis is
is usually dominated by species in the Myrtaceae. present in some tropical botanic gardens.
The altitudinal range of A. borneensis is substan-
tial, from near sea level to ca. 2400 m a.s.l., but with
greater abuncance below ca. 1200 m a.s.l. Agathis dammara (Lamb.) Rich. & A. Rich., in
A. Richard (ed.) Comm. Bot. Conif. Cycad.: 83.
Conservation 1826. Pinus dammara Lamb., Descr. Pinus 1: 61,
t. 38. 1803. Type: Illustration in Rumphius, Herb.
This species has been very heavily over-exploited in Amboinense 2: 174, t. 57. 1741 (lectotype).
many areas and as a result its total area of occupancy
(AOO) is estimated to have at least been reduced by Agathis celebica (Koord.) Warb., Monsunia 1: 185.
half and this is still ongoing. Stands covering an esti- 1900; Dammara celebica Koord., Meded. Lands
mated total of 30,000 ha discovered in Kalimantan Plantentuin 19: 263. 1898.
in the 1930s had effectively been logged out by the Agathis philippinensis Warb., Monsunia 1: 185. 1900.
mid 1960s. Most stands outside the few well pro-
tected nature reserves (mostly situated in the Malay Etymology
Peninsula and in Sabah) have been seriously depleted
and it is doubted that regeneration will be sufficient Dammar is the local (Moluccan) name for the resin,
to restore the losses. Habitat degradation has caused both subfossil (copal) and recent, of this tree.
further reductions in recruitment of young trees to
replace felled ones. Vernacular names
IUCN: EN (A4cd)
Amboina pitch tree; dammar raja (Indonesia); dam-
Uses mar malolo, dammar lulu (Sulawesi); almaciga,
saleng (Philippines); kalne, kssi, oenela (Maluku
This species is one of the most valuable and sought [Moluccas]) and many other local names (see e.g. in
after timber trees in Southeast Asia. It produces Flora Malesiana, ser. 1, 10 (3): 438, 1988).
lightweight, almost white to pale yellowish wood
with no visible growth rings and a very fine and even Description
texture and without resin. It is not durable, so it will
mostly find indoor uses, but these are many, from Trees to 55(65) m tall and 3.54 m or more d.b.h.
light construction and carpentry, joinery, masts for with clear cylindrical bole up to 2025 m and a
broad crown radiating above. Bark variable; smooth (Whitmore, 1980) were by De Laubenfels separated
or dippled and lenticellate or rough and scaly, colour as distinct species. De Laubenfels (1988) complicated
grey, dark brown or blackish outside, reddish brown the nomenclatural knot by asserting that, under the
or yellowish brown under outer, exfoliating layers. rules of ICBN and in case a proposal to reject the
Leaves subopposite, thick, multinerved, coriaceous, name Pinus dammara Lamb. in favour of A. borneen-
glabrous, light green. Leaves on saplings and young sis Warb. was not accepted at the next International
trees and/or in shade lanceolate to (narrowly) ellip- Botanical Congress, A. borneensis would have to
tic, up to 14 cm long, 1.54 cm wide, often acute, be called A. dammara. His taxonomic views were
sometimes acuminate; leaves in crowns of mature followed in the World Checklist & Bibliography of
trees distinctly petiolate; shape variable, ovate to Conifers (Farjon, 1998, [2001]). This proposal was
156 ovate-elliptic or sometimes (narrowly) lanceo- indeed rejected, but only if one sinks A. borneensis
late, 2.58(9) cm long, (1)24 cm wide, with an (the western species) taxonomically into A. dam-
obtuse or acute apex. Pollen cones axillary, solitary mara (the eastern species) would the resulting taxon
on a 05 mm long, stout peduncle, when imma- have to bear that name. The type of A. dammara is
ture a small cylinder, 12 cm long, 68 mm diam., from Amboina, not from Borneo, and the pollen
elongating past anthesis to 34 cm long and up to cones of the two are very different. These are sepa-
12 mm wide, with 48 decussate (in 24 pairs) 24 rate species. This is not so with the trio A. celebica,
mm wide, free spreading bract scales at their base. A. philippinensis and A. dammara; the distinctions
Microsporophylls in imbricate arrangement; head mentioned in Flora Malesiana (De Laubenfels, 1988)
slightly convex towards the upper, rounded or retuse, e.g. acuminate juvenile leaves (meaning: leaves on
minutely erose-denticulate margin, 22.5 mm wide juvenile trees) are found occasionally in specimens
and 11.5 mm high in mature cones, bearing 36 from all (major) Malesian islands and are clearly
oblong pollen sacs. Seed cones solitary on thick just one of the possible shapes of these highly vari-
peduncles, broadly ellipsoid to globose, to 1013 cm able leaves. The diagnostic pollen cones are a much
diam., smooth, green, resinous, ripening brown. better organ than the leaves on young trees to look
Cone scales with slightly rounded, incurved upper for consistent characters, and they unite A. celebica,
margins, ca. 3.5 cm long, 3.54.5 cm wide in larger A. dammara and A. philippinensis, a species which
cones, roughly triangular with rounded corners and therefore bears the earliest name. Other characters,
more or less flanged on either side. Seeds 1215 78 some involving leaves of mature trees, do separate
mm, ovoid-oblong, with two unequal wings; largest some of the montane populations in Borneo, and are
wing ca. 20 13 mm; smallest wing a small blunt here not sunk into A. dammara.
triangle 35 mm wide opposite largest wing.
Distribution
Taxonomic notes
Malesia: Maluku [Moluccas], Philippines, Sulawesi.
The nomenclature of this species is complicated, TDWG codes: 42 MOL PHI SUL
reflecting in part conflicting views on its taxonomy
for nearly 200 years. In more recent time, Whitmore Ecology
(1980) included not only all trees belonging to the
genus occurring in Sulawesi, the Moluccas and the Agathis dammara occurs in lowland to upland tropi-
Philippines, but also several montane populations in cal rainforest as scattered emergent trees. In lowland
Borneo and the Malay Peninsula in A. dammara. De to lower montane rainforest it can be associated with
Laubenfels (1988) considered A. philippinensis a dis- Dipterocarpaceae and/or Fagaceae; however, Agathis
tinct species, occurring in the Philippines but also often retreats to ridges with thin, rocky soils or to
in Sulawesi and on the Moluccas. He furthermore water-logged areas where these dominant angio-
recognized A. celebica in Sulawesi and the Moluccas, sperms are less vigorous. The species occurs on a
with a few outliers in the Philippines. In addi- wide variety of substrates, from white sand to peaty
tion, some of the montane populations cited above soils, volcanic soils, metamorphic rock such as ser-
pentine or schist, or limestone. The altitudinal range Agathis flavescens Ridl., Kew Bull. 1914: 332. 1914.
of A. dammara is from near sea level to ca. 2200 m Agathis dammara (Lamb.) Rich. & A. Rich. subsp.
a.s.l., but with greater abuncance below ca. 1200 m flavescens (Ridl.) Whitmore, Pl. Syst. Evol. 135 (12):
a.s.l. 59. 1980; Agathis celebica (Koord.) Warb. subsp.
flavescens (Ridl.) Veldkamp & Whitmore, Taxon 33
Conservation (2): 346. 1984. Type: Malaysia: Peninsular Malaysia,
Pahang, Gunung Tahan, H. N. Ridley 16023
This species has been over-exploited in many areas (holotype K).
and as a result its total area of occupancy (AOO)
is estimated to have at least been reduced by 30% Etymology
or more and this is still ongoing. The tapping of 157
resin when exploited too intensively has killed The species epithet (Latin flavescens = yellowish or
large numbers of trees in the forests, especially in pale yellow) refers to leaf colour.
the Philippines. Habitat degradation has caused
further reductions in recruitment of young trees Vernacular names
to replace felled ones. There is now a total ban on
cutting Agathis trees in the remaining forests in the Tahan agathis.
Philippines, but there is still illegal logging going on
in some areas. Description
IUCN: VU (A4cd)
Trees to 18(21) m tall and 1.5 m or more d.b.h. with
Uses short cylindrical bole and a spreading, rounded or
more or less flat-topped crown. Bark smooth or
Large trees of this species are highly valuable tim- dippled and lenticellate, colour grey, reddish brown
ber trees, yielding large sizes of straight, knot-free, under outer, exfoliating layers. Leaves subopposite,
strong and light coloured sawn timber. It is used thick, multinerved, coriaceous, glabrous, (yellow-
for construction as beams, joists and frames, in car- ish?) green. Leaves on saplings and young trees and/
pentry for joiners work, for boat building includ- or in shade ovate-lanceolate, up to 8 cm long and
ing oars due to its elasticity, idem for light aircraft 3 cm wide, acute or slightly acuminate; leaves in
as a substitute for Sitka spruce, and for floor boards, crowns of mature trees distinctly petiolate, ovate to
panelling and furniture as well as picture frames, ovate-lanceolate, (2)35(7) cm long, (1)23 cm
pencils, rulers and T-squares. Lower grade wood is wide, with an obtuse or rounded apex. Pollen cones
used as pulpwood in the paper industry. In parts of axillary, solitary on a (2)510(15) mm long, stout
Sulawesi where this tree no longer occurs subfossil peduncle, when immature a small cylinder, 12 cm
resin (copal) is found and mined, most likely pro- long, 68 mm diam., elongating past anthesis to 34
duced by trees that lived there many centuries ago. cm long and up to 10 mm wide, with 4 decussate (in
This hardened resin is the source for products like 2 pairs) 24 mm wide, free spreading bract scales
paints and varnishes and is searched for and dug up at their base, subtended by two much longer (up to
by copal diggers as itinerant labourers setting up 30 10 mm) leaf-like bracts. Microsporophylls in
camps in the forest, often following logging opera- imbricate arrangement; head convex towards the
tions. This resin was formerly a much more impor- upper, rounded or retuse, minutely erose-denticu-
tant product of this tree to the local inhabitants late margin, 2 mm wide and 1.5 mm high in mature
than its wood. Agathis dammara is used in forestry cones, bearing 36 oblong pollen sacs. Seed cones
plantations, mainly in Jawa, where the genus does solitary on thick peduncles, broadly ellipsoid to
not occur naturally. It is very rare in tropical botanic globose, to 78 cm diam., with bossed scales (sur-
gardens, where trees labeled A. dammara may be A. face not smooth), green, resinous, ripening brown.
borneensis instead. Cone scales with rounded to angular or bossed,
thick upper margins, ca. 3 cm long, 4 cm wide in which is mostly covered in heath-like dwarf shrubs.
middle part of full-grown cones, rounded triangu- Agathis borneensis occupies a lower zone on this
lar to broadly reniform and more or less flanged on mountain covered with taller forest; the two appear
either side. Seeds 1113 78 mm, ovoid and flat- to be separated by a belt of vegetation devoid of
tened, with two unequal wings; largest wing ca. 15 Agathis (Chung-Lu Lim, FRIM unpublished data).
10 mm; smallest wing a small blunt triangle 3 mm
wide opposite largest wing. Conservation
Distribution Uses
Fiji: Kadavu Is., Viti Levu, Vanua Levu; Solomon The wood of this species is white or sometimes with
Islands: Santa Cruz Group (Utupua Is., Vanikoro a reddish hue and known in Fiji as Dakua wood and
Is.); Vanuatu: Anatom Is., Erromango Is., Tanna Is. in the Santa Cruz Group as Vanikoro kauri. It is very
TDWG codes: FIJ SCZ VAN valuable and used for construction, for flooring in
164 houses, for masts, booms and spars in sailing boats,
Ecology for carpentry and for furniture making. The resin
exuded from the bark is fragrant and inflammable
Agathis macrophylla is an emergent tree in lowland and is (was) burnt to provide light. Recent resin is
to low montane tropical rainforest; usually growing transparent and nearly colourless but weathers white
in soils derived from volcanic rocks like basalt. Its in contact with air and sunlight; subfossil resin has
altitudinal range is recorded from herbarium col- a yellowish or orange-brown hue approaching
lections as being between 75 m and 900 m a.s.l. In some types of amber, which is completely fossilized
Fiji on the main islands it is most common between (matured) resin from conifers. Resin is tapped from
600 m and 900 m a.s.l. The species was the subject trees, but also dug from the ground (subfossil resin)
of a pioneering study on its role in the rain forests and used in making varnishes, pottery glazing, and
of the type locality, which demonstrated that unlike dying cloth black with the smoke from burning it.
Agathis australis in New Zealand, and many other Fijian kauri pine has been planted as a forestry tree
species which exhibit a regeneration strategy based in the Solomon Islands (Santa Cruz Group) and
on periodic landscape-scale disturbance (cf. Enright elsewhere in the SW Pacific in an attempt to obtain
& Hill, 1995) A. macrophylla appears to behave as timber more sustainably from a truly renewable
a normal component of rainforests dominated by resource. It is also in cultivation in some green-
angiosperms. This means that it is capable of small- houses of botanic gardens.
gap regeneration like other large forest trees.
Agathis microstachya J. F. Bailey & C. T. White,
Conservation Contr. Queensland Fl. Bot. Bull. 18: 13. 1916. Type:
Australia: Queensland, Cook District, H. W.
Overall, this species was listed as Near Threatened Mocatta s.n. (holotype BRI). Fig. 35
(IUCN Redlist 1999). However, individual island
populations may well be severely threatened, e.g. Etymology
that on Utupua in the Santa Cruz Group. Doyle
(in Farjon & Page, 1999) assessed the species sepa- The species epithet derives from the Greek micros =
rately for each of the island groups as follows: Fiji: small and stachys = ear of corn (maize, not known in
VU Vulnerable, IUCN 1994-criteria A2d (>20% ancient Greece) or a flower spike, and alludes to the
reduction within next three generations based small male strobili.
on current exploitation levels) and B2e (continu-
ing decline in the number of mature individuals). Vernacular names
Santa Cruz Islands: NE not evaluated. Vanuatu:
VU Vulnerable, IUCN 1994-criteria A2d and Bull kauri, Bull pine, Atherton kauri pine
B2e. Agathis macrophylla is now quite rare outside
plantations in the Santa Cruz Group, though an Description
unlogged population apparently survives on the
upper Lawrence River on Vanikoro. A re-assessment Trees to 50 m tall and 2.5 m or more d.b.h. with
using version 3.1 of the IUCN criteria (IUCN, 2000) clear cylindrical bole to 35 m and a rounded or open
crown with spreading to ascending branches. Bark much larger than the smallish seed cones of A. atro-
smooth or sometimes scaly, exfoliating in irregu- purpurea. There is a general leaf shape difference in
lar coarse scales of up to 15 cm across; outer bark leaves on mature trees in that A. microstachya has
brown or grey-brown, with bluish purple patches obtuse or rounded leaf apices, while A. atropurpurea
under newly fallen scales; inner bark reddish, exud- has only obtuse apices, but this character is less
ing white resin. Leaves subopposite, thin or thick, reliable.
multinerved, coriaceous, glabrous, light green.
Leaves on saplings and young trees and/or in shade Distribution
broadly lanceolate to elliptic, 59 cm long, 1.53.5
cm wide; leaves in crowns of mature trees lanceo- Australia: NE Queensland (Cook District).
late to elliptic, 38 cm long, (0.8)12.2 cm wide, TDWG codes: 50 QLD-QU 165
short petiolate and with an obtuse or rounded apex.
Pollen cones axillary, solitary on a 01 mm long Ecology
peduncle, subglobose becoming short cylindrical,
when full grown 11.6 cm long, 58 mm diam., with Agathis microstachya occurs mainly scattered in low-
810 decussate (in 45 pairs), imbricate, 12.5 mm land to low montane semi-evergreen tropical rain-
wide bract scales at their base. Microsporophylls in forest in mountains E of the Atherton Tablelands,
tesselate arrangement; heads prismatic, with raised but also less frequently on these plateaus. Its alti-
central part irregularly pentagonal or hexagonal and tudinal range is between 400 m and 1100 m a.s.l.
becoming free at anthesis, 0.5 mm wide, 0.50.7 mm It is an emergent species in a species-rich canopy
high in mature cones, bearing 25 pollen sacs. Seed of angiosperm trees. Soils are loamy sands or light
cones solitary on thick peduncles, globose or ovoid, clays and derived from acidic to neutral silicate-rich
7.511.5 cm long, 6.510 cm wide, smooth or nearly rock, usually of volcanic origin. Annual rainfall var-
so, green or slightly glaucous, ripening brown. Cone ies, with maxima to over 3000 mm on ocean-facing
scales with thick, rounded, incurved upper margins, mountain slopes.
2.53.5 cm long, 3.34.5 cm wide, broad triangular
with rounded corners and more or less flanged on Conservation
either side. Seeds 1012 68 mm, ovoid, flattened,
with two unequal wings; largest wing 2025 1013 Before 1985 the population of A. microstachya had
mm; smallest wing a more or less triangular point been nearly halved by logging but 70% of the for-
25 mm long, opposite largest wing. ests are now protected. The remaining population
is estimated to consist of fewer than 10,000 mature
Taxonomic notes trees. Under IUCN Red List criteria (version 3.1,
2001) this species would therefore qualify for the
Agathis atropurpurea and A. microstachya are more status VU, but due to cessation of large-scale logging
or less sympatric, but remain often separated from and effective protection in a National Park and other
each other by altitudinal range (with A. micro- protected areas, its threat with extinction has been
stachya usually occurring at a lower altitude), and greatly reduced. Logging continues in unprotected
slight habitat differences, e.g. soil conditions, which areas, but the decline appears to have been halted.
are generally poorer with A. atropurpurea. They do IUCN: NT
not seem to hybridize. The most consistent differ-
ence between them is the morphology of the micro- Uses
sporophylls in the pollen cones (these cones are
small in both species). In A. atropurpurea, these are Bull kauri is a valuable timber tree and was formerly
imbricately arranged (the most common situation in intensively logged and taken to local sawmills. The
the genus), while the microsporophylls in A. micro- timber is soft, light, easy to work and polishes well.
stachya are tesselate, i.e. laying together like paving It is used for house framing, flooring, and joinery,
stones, with no parts overlapping. The shapes of the as well as veneer production. This species is planted
microsporophyll heads are also markedly different. on a limited scale mainly in arboreta and tree collec-
The seed cones of A. microstachya are, when mature, tions in parks in Queensland.
Agathis montana de Laub., Trav. Lab. Forest. Distribution
Toulouse T. 1 (8, 5): 2. 1969. Salisburyodendron
montana (de Laub.) A. V. Bobrov & Melikyan, New Caledonia (Massif du Pani, Roches d
Komarovia 4: 63. 2006. Type: New Caledonia: Ouaime).
Grande Terre, Province Nord, Mt.Pani, M. Schmid TDWG codes: 60 NWC
1420 (holotype P).
Ecology
Etymology
Agathis montana is abundant to dominant from
The species epithet means from the mountain and 11001200 m upwards on the Pani massif, the
166 refers to its habitat. highest and wettest mountains in New Caledonia.
It occurs in fern-rich rain forest habitat with
Vernacular names Retrophyllum comptonii, the only other large tree, on
reddish soil overlying micaschists. On the summit
No common names are known for this species. of Mt. Pani Agathis montana occurs together with
Araucaria schmidii on or near exposed outcrops of
Description rock and steep summit slopes, where it becomes
stunted. The altitudinal range of this species is from
Medium size trees to 1520 m tall, either multi- 950 m a.s.l. to 1600 m a.s.l. on the summit crest.
stemmed from near base or with clean bole for 810
m; sometimes almost flat-topped in appearance but Conservation
crown usually very sparse, with long, spreading,
crooked branches. Young trees possibly conform- IUCN: NT
ing to Rauhs architectural model, with whorled
tiers of assurgent branches forming a conical crown. Uses
Bark reddish brown or tan to grey with numerous
small, granular lenticels, flaking, coming away from No uses are recorded of this species. The first botani-
the tree in irregular small pieces, excuding copious cal collection is from as recently as 1939 and the alti-
white resin. Leaves subopposite, thick, multinerved, tude and relative inaccessibility of all the known
coriaceous, glabrous, glaucous on the abaxial sur- stands may have prevented earlier utilization, e.g.
face. Leaves of saplings and young trees broadly the collecting of resin. The commonly crooked shape
ovate, 811 cm long, 2.43.8 cm wide; leaves in crown and shortness of the boles render it less suitable for
of mature trees becoming more lenticular, 5.59 cm timber than most other species in the genus. It is not
long, 1.42.2 cm wide. Pollen cones axillary, solitary, known to be in cultivation.
short pedunculate or sessile, with a tight basal-bract
cluster of 58 imbricate, 12 mm wide bract pairs and
without a pair of leaf-like bracts, cylindrical, 45 cm Agathis moorei (Lindl.) Mast., J. Roy. Hort. Soc.
long but expanding to 7 cm or more past anthesis, London 14: 197. 1892. Dammara moorei Lindl.,
810 mm wide. Microsporophylls imbricate; micro- J. Hort. Soc. London 6: 271. 1851; Salisburyodendron
sporophyll heads umbonate, apices weakly acumi- moorei (Lindl.) A. V. Bobrov & Melikyan,
nate, 2.02.6 mm wide and 2.22.5 mm high. Seed Komarovia 4: 63. 2006. Type: New Caledonia:
cones axillary, solitary on stout peduncle, globose to Grande Terre, [on East Coast], C. Moore s.n.
obovoid, ca. 9 cm long, 7 cm wide, smooth, green to (holotype CGE).
glaucous green ripening brown, very resinous. Cone
scales with rounded, thick and slightly incurved Agathis corbassonii de Laub., Trav. Lab. Forest.
upper margins and thin, fragile lateral margins, ca. Toulouse T. 1 (8, 5): 2. 1969. Salisburyodendron
3 cm long and wide. Seeds ovoid, 68 5 mm, with corbassonii (de Laub.) A. V. Bobrov & Melikyan,
one large ovate wing and a small, triangular wing on Komarovia 4: 63. 2006.
the opposite side (no intact wings seen).
Etymology Taxonomic notes
This species was named after Charles Moore (1820 Agathis corbassonii was segregated from A. moorei
1905), a Director of the Royal Botanic Gardens on the basis of its very reddish brown bark and its
Sydney, who collected the type specimen. narrow leaves which are often glaucous beneath.
The possibility of separate species (kaori blanc, kaori
Vernacular names rouge) was first suggested by Michel Corbasson,
Director of the Tropical Forestry Institute in Noumea.
Moores kauri; kaori blanc, kaori rouge (French in However, the variation in leaf sizes and shapes in A.
New Caledonia) moorei is perhaps greater than in any other species
in the SW Pacific (T. G. Waters, unpublished D.Phil 167
Description thesis, Oxford 2008) and those specimens ascribed to
A. corbassonii appear to fall mostly within the ranges
Trees to 40(50) m tall and 1 m or more d.b.h. with observed in A. moorei. Glaucousness is not a sound
clear cylindrical bole to 25 m; crown conical or character for taxonomic distinction as it so often var-
rounded in trees of medium size, becoming more ies within populations in conifers. Field observation
open and irregular in old trees. Bark smooth or strongly indicates that shaded boles within forests
rough and scaly, peeling in small or large, irregular have reddish brown bark, while sun-exposed trees
flakes; colour variable in forest reddish brown to develop grey bark through a weathering process.
tan, but becoming greyer (and harder) with expo- This exposed bark also becomes harder than on trees
sure to sun, often resinous. Branches spreading. in moist forests. [It is understandable that local for-
Leaves subopposite, thick, multinerved, coriaceous, esters took note of these bark differences, which is
glabrous, light green, sometimes glaucous on the often all one can see of a large tree in tropical rain-
abaxial (lower) side. Leaves on saplings and young forest.] The number of bract scales at the base of the
trees and/or in shade lanceolate to elliptic, 1020 cm pollen cones in A. moorei is not consistently 16 (8
long, 24 cm wide; leaves in crowns of mature trees pairs in decussate arrangement) as is stated in some
narrowly ovate-elliptic, 4.57 cm long, (6)812 mm published descriptions, but can be fewer; it is a maxi-
wide with an obtuse apex. Pollen cones axillary, mum number distinguishing the pollen cones of A.
solitary on a slender, (1)812 mm long peduncle, moorei from other species in New Caledonia. Agathis
cylindrical, when full grown 2.53(5) cm long, corbassonii is here treated as a synonym of A. moorei.
69 mm diam., with 816 decussate (in 48 pairs), The lower number of bract scales recorded for A. cor-
imbricate, 45 mm wide bract scales at their base. bassonii (based on few good specimens) is therefore
Microsporophylls in imbricate arrangement; head included in the character states as found in A. moorei,
more or less angular but nearly flat, 13 mm wide, and in agreement with the measurements compiled
0.752 mm high in mature cones; margin minutely by Waters and his judgement on the matter.
erose-denticulate. Seed cones solitary on thick
peduncles, subglobose to globose, 1015 cm long, Distribution
912 cm diam., smooth, green or glaucous green,
(very) resinous, ripening brown. Cone scales with New Caledonia: Grande Terre.
slightly rounded, incurved upper margins, 34 cm TDWG codes: 60 NWC
long, 34 cm wide, roughly triangular with rounded
corners and more or less flanged on either side. Seeds Ecology
1520 810 mm, ovoid-oblong, with two unequal
wings; largest wing 2030 1520 mm; smallest Agathis moorei occurs in dense rain forests on soil
wing reduced to a narrow appendix 35 mm long, derived from schists, micaschists, gneiss, and ser-
opposite large wing. pentines. It is usually restriced to non-ultramafic
soils, either forming small groves of pure Agathis ating above. Bark sparsely lenticellate, hard and
moorei, or scattered through mixed angiosperm for- smooth, exfoliating in small plates to 4 mm thick,
est. Its altitudinal range is from (30) 100 m to 700 colour dark brown or greenish purple outside, red-
(1000) m a.s.l. dish brown under outer, exfoliating layers, exuding
light yellow resin. Leaves subopposite, thick, multi-
Conservation nerved, coriaceous, glabrous, light green and often
glaucous underneath. Leaves on saplings and young
This species has been heavily exploited for its tim- trees and/or in shade broadly lanceolate, 47 cm
ber in the past and consequently it has disappeared long, 23.5 cm wide, acute or slightly acuminate;
or become very scarce in many parts of the for- leaves in crowns of mature trees distinctly petiolate,
168 est, especially in more accessible localities. Even in oval or ovate to nearly orbiculate, (2)2.54(6) cm
pre-European times its wood was preferred for the long, 12.5 cm wide, acutish or obtuse to rounded
making of outrigger canoes. Substantial decline has at apex. Pollen cones axillary, solitary on a 23 mm
occurred also within recent years and may continue long peduncle, when immature from subglobose
because illegal logging is a particular concern (Watt becoming nearly cylindrical, 68 mm long, 45 mm
in Farjon & Page, 1999). However, the species still diam., only slightly elongating past anthesis to 1215
occurs over much of the island of Grande Terre and, mm long and 56 mm wide, with 46 decussate (in
given adequate protection, could recover in many 23 pairs) 23 mm wide, free spreading bract scales
areas to former abundance. at their base. Microsporophylls in imbricate arrange-
IUCN: VU [B1ab(iii)+2ab(iii). C1] ment; head slightly convex towards the upper,
rounded to angular, minutely erose-denticulate
Uses margin, 1.21.5 mm wide and 1 mm high in mature
cones, bearing 24 oblong pollen sacs. Seed cones
Moores kauri is a valuable timber tree and its wood is solitary on thick peduncles, ovoid to subglobose,
(was) much used for carpentry, joinery, veneer, and to 8 cm long and 6 cm diam., more or less rough
for the construction of pirogues (outrigger canoes). with raised seed scale margins especially in imma-
In the extreme southern part of Grande Terre some ture cones, green, resinous, ripening brown. Cone
limited forestry plantations of this species have been scales with more or less rounded, bossed, thick and
established on ultramafic soils. strongly incurved upper margins, ca. 2 cm long and
3 cm wide in mature cones, flabellate to nearly reni-
form with rounded corners and more or less flanged
Agathis orbicula de Laub., Blumea 25 (2): 540. on either side. Seeds ca. 10 6 mm, ovoid, flattened,
1979. Type: Malaysia: Sarawak, 5th Division, Lawas, with two unequal wings; largest wing ca. 12 6 mm;
[Bumbong Rumah, N of Lawas], D. J. Laubenfels smallest wing a small triangle of 2 mm opposite large
de P 614 (holotype L). wing, or absent.
The species epithet (Latin orbis = globe) describes Agathis orbicula has similar shapes and sizes of
the shape of (some of) the leaves as being nearly leaves to A. kinabaluensis, but those of the latter are
round. not glaucous on the abaxial (lower) side. The pol-
len cones of A. orbicula are among the smallest in
Vernacular names the genus (after anthesis) and lack the two elongated
basal bracts observed in A. kinabaluensis. Agathis
tumuh (Murut); tubu (Kenyah); bulok (Iban) orbicula occurs at lower altitudes in tall forest and
consequently is a tall tree; A. kinabaluensis occurs at
Trees to 40 m tall and 2 m or more d.b.h., with clear high montane altitudes and is mostly a short, stunted
cylindrical bole up to 20 m and a broad crown radi- tree, but grows tall when in high forest.
Distribution Agathis ovata (C. Moore ex Vieill.) Warb.,
Monsunia 1: 186. 1900. Dammara ovata C. Moore
Borneo: Indonesia (Kalimantan Timur); Malaysia ex Vieill., Ann. Sci. Nat. Bot., sr. 4, 16: 56. 1862;
(Sabah, Sarawak). Salisburyodendron ovata (Vieill.) A. V. Bobrov
TDWG codes: 42 BOR-KA BOR-SB BOR-SR & Melikyan, Komarovia 4: 63. 2006. Type: New
Caledonia: Grande Terre, Province Sud, Col de
Ecology Yat, [Monts dUnia], E. Vieillard 1263 (holotype
P). Fig. 36, 37
Agathis orbicula occurs at lower altitudes than A.
lenticula in evergreen tropical rainforest and high Dammara hypoleuca C. Moore ex Henkel & W.
kerangas. It is, like that and other species at these Hochst., Syn. Nadelhlz.: 217. 1865; Agathis hypo- 169
lower altitudes (for this species between 450 m and leuca (C. Moore ex Henkel & W. Hochst.) Warb.,
1050 m a.s.l.) a tall emergent tree. Monsunia 1: 186. 1900; Salisburyodendron ovata
(Vieill.) A. V. Bobrov & Melikyan subsp. hypoleuca
Conservation (C. Moore ex Henkel & W. Hochst.) A. V. Bobrov &
Melikyan, Komarovia 4: 63. 2006.
This species is known from seven localities. In addi-
tion to those mapped in Flora Malesiana, ser. 1, 10 Etymology
(3): 437, f. 78 ( De Laubenfels, 1988) there are two
in Kalimantan Timur. Exploitation of all tall trees The species epithet refers to the often ovate leaves.
of Agathis in Borneo has been intensive and is still
ongoing. As a consequence, we can infer a decline Vernacular names
even though this species is not distinguished by for-
esters or loggers from the more widespread species Scrub kauri; kaori de montagne, kaori nain (French
A. borneensis, which occurs in the same general area. in New Caledonia)
The true rate of decline could be impossible to esti-
mate, as logged trees and their stumps are no lon- Description
ger identifiable to species and cut trees assumed to
belong to the more common A. borneensis may have Small, often shrubby trees up to 10 m tall (occa-
been A. orbicula. It may have been present in several sionally to 25 m in forest), usually with a short
more than the presently known localities. trunk to 80 cm d.b.h. and branching from near the
IUCN: VU [B2ab (ii, iii, v)] base, with orthotropic primary branches (Rauhs
model) forming a flat-topped or irregular, spread-
Uses ing crown. Leaf-bearing branches and cones held
in dense pseudo-verticillate, upright or ascending
The wood of this species has the same properties as clusters at tips of main branches, often glaucous to
other lowland species of Agathis in Borneo and is farinose. Bark with deep fissures, hard, slowly flak-
used for the same purposes; see therefore under A. ing off and forming irregular polygonal patches and
borneensis. The resin is said to have a yellow colour, scales on main trunks and branches, tan or reddish
but since the resin of A. lenticula and other species soon becoming grey. Leaves subopposite, thick,
comes out clear or white and tends to turn yellowish multinerved, coriaceous, glabrous, light green or
on exposure to air, this may be a somewhat doubt- glaucous green on the abaxial (under) side. Leaves
ful property and is not known to be relevant to its on saplings or young trees well spaced, spreading in
applications. This species is not known to be in cul- a plane, obovate-truncate, 46 cm long, 1015 mm
tivation, but since plantations for forestry have not wide. Leaves in crowns of mature trees crowded,
made the taxonomic distinction between this spe- often in 4 more or less erect rows, similar in shape,
cies and A. borneensis, it may be present in some often somewhat wider and with the leaf apex trun-
plantations. cate or notched, 48.5 cm long and 1.55 cm wide
but occasionally almost orbiculate in shape. Pollen Conservation
cones axillary, solitary on a stout 420 mm long
peduncle, more or less fusiform (widest in the mid- This species is widespread but scattered in ultramafic
dle) but with an obtuse apex, 35 cm long, 1015 mm maquis in Province Sud, and protected in some
diam.; subtended by two small leaf-like bracts and major reserves. Fire remains a threat in many areas
45 pairs of imbricate, short, rounded bract scales; where its incidence has been increased by humans,
basal-bract cluster usually narrower than widest and a continuing decline is projected to occur.
point of pollen cone. Microsporophylls imbricate, IUCN: EN [B1 ab(iii)+2ab(ii, iii)]
microsporophyll head 24.5 mm wide, 1.754.5 mm
high in mature cone, more or less convex with entire Uses
170 margin. Seed cones globose, ca. 6 cm long, 5 cm
wide; seed scales bossed to shortly rostrate: mature Due to its usually low stature and crooked habit,
cones not smoothly globose. Cone scales 1620 mm this species is not exploited for timber, although an
long, 2226 mm wide, with a rounded, thick, more occasional tall specimen may have been logged from
or less shortly rostrate (bossed) upper margin and forest habitat. It is not known to be in cultivation
thin, rounded lateral margins ending in a cuneate outside a few collections in botanic gardens in the
base. Seeds ovoid, 911 mm long, 78 mm wide, (sub-)tropics.
with truncate base and apex; wings of unequal size;
largest wing 1520 mm long and 912 mm wide and
extended at a right angle to the seed; smaller wing Agathis robusta (C. Moore ex F. Muell.) F. M.
opposite with a rounded edge, ca. 3 mm long and 5 Bailey, Syn. Queensland Fl.: 498. 1883.
mm wide.
Etymology
Distribution
The species epithet robust may refer to the leaves,
New Caledonia: Grande Terre, mainly in Province which are often larger than those of the few species
Sud (Massif du Sud) but with two localities fur- known in 1857 when C. Moore coined the name.
ther north, one as far as Commune de Houalou in
Province Nord. Vernacular names
TDWG codes: 60 NWC
Queensland kauri pine, Smooth-barked kauri;
Ecology asong, muwaka, ogapa (Papua New Guinea, mostly
for subsp. nesophila).
Agathis ovata is usually growing on ultramafic soils in
both maquis and (more rarely) forest environments. Its Description
altitudinal range is from 30 m to 1050 m a.s.l. In maquis
environments it commonly occurs with Araucaria Trees to 50 m tall and 2 m or more d.b.h. with clear
spp., Babingtonia leratii, Xanthostemon spp., Grevillea cylindrical bole to 30 m and a rounded or open crown
spp., various members of the Cyperaceae, Pteridium with spreading to ascending branches. Bark smooth,
esculentum, Dracophyllum spp., various members often lenticellate, exfoliating in roundish small scales
of the Cunoniaceae, Myodocarpus spp., Dacrydium of up to 5 cm across; outer bark brown, reddish under
araucarioides, Lomandra insularis, Gymnostoma spp., newly fallen scales; inner bark pinkish, exuding
Styphelia spp., Asplenium spp., Hibbertia spp., and white resin. Leaves subopposite, thin or thick, mul-
lichens such as Cladia retipora. In forest environ- tinerved, coriaceous, glabrous, light or mid lustrous
ments it occurs in mixed tropical rain forest, usually green above, dull green below. Leaves on saplings
on drier, exposed slopes or ridges, but including for- and young trees and/or in shade broadly lanceolate,
ests otherwise dominated by species of Nothofoagus. 812(14) cm long, 35 cm wide, sometimes acu-
Here it may occasionally attain 2025 m and, due to minate; leaves in crowns of mature trees narrower,
competition with other trees, grow as an upright tree. lanceolate, oblanceolate or elliptic, 511(14) cm
long, 13 cm wide, distinctly petiolate and with an of nutrient-poor soils derived from granite or other
obtuse or rounded apex. Pollen cones axillary, soli- igneous rock, including ultrabasic types.
tary on a 210(18) mm long, stout peduncle, cylin-
drical, when full grown 46(7) cm long, 813 mm Uses
diam., becoming flexible, with 810 decussate (in
45 pairs), imbricate but spreading, 36 mm wide The Queensland kauri pine is a valuable timber tree
bract scales at their base. Microsporophylls in tes- and was exploited in the past in Queensland, but
selate (or very weakly imbricate) arrangement; logging of natural stands has all but ceased there. In
head obtuse-triangular to rhombic in outline, with New Guinea the timber of this species is used locally
a raised, prismatic, nearly flat or keeled central part for house construction and the resin is tapped for
12 mm wide, 0.71.5 mm high in mature cones; various uses, but since this tree occurs very scat- 171
margins partly hidden, bearing 28 pollen sacs. tered in the forest it is not of economic impor-
Seed cones solitary on thick peduncles, subglobose tance to the timber industry. Export of round logs
or obovoid, 815 cm long, 810 cm wide, smooth, of Agathis from Papua New Guinea is banned. This
green and resinous, ripening brown. Cone scales species is widely planted as an ornamental in the
with slightly rounded, incurved upper margins, ca. tropics and subtropics and is present in most (sub)
3 cm long, 3.54.2 cm wide, roughly triangular with tropical botanic gardens and many parks. Most of
rounded corners and more or less flanged on either the provenance of these trees will be populations
side. Seeds 1214 78 mm, ovoid, flattened, with in Queensland, especially from the northern pop-
two unequal wings; largest wing 2025 1215 mm; ulation in Cook and Kennedy Districts. It is also
smallest wing a small acute triangular point 34 mm planted locally with more utilitarian aims, especially
long, opposite largest wing. resin tapping in PNG.
Latin: amentum = string; referring to the racemose The colour of the stomatal bands is more or less
arrangement of pollen cones; Taxus is the classical white in all species, but will turn brownish in some
Latin name for yews. when leaves are older than one year.
Shrubs or small trees to 7 m tall. Bark smooth, exfoliat- Amentotaxus argotaenia is widespread and occurs on
ing in thin flakes. Branches few, spreading or ascend- limestone as well as sandstone, shale or granite, and
ing. Foliage branchlets opposite, spreading at 2570 in ravines, on steep slopes or cliffs, on summits and
from the leading shoot axis, ascending or spreading, ridges and in mountain forests along shady stream
subterete, with alternatingly twisted grooves running banks. The altitudinal range is between 600 m and
from one leaf base to the next above, green in the first 1100 m a.s.l. Associated trees and shrubs vary with
year, turning from greenish yellow to orange-brown the types of rock, mainly between limestone and
in following years, terminating in a conical bud with other rocks. On limestone, it may grow with Pinus
ovate-triangular, acute scales. Leaves spreading dis- kwangtungensis, Nageia spp., Podocarpus neriifolius, 175
tichously at 4590(95) from shoot axis, linear or P. pilgeri, P. macrophyllus and Taxus chinensis. On
linear-lanceolate, (2)39(11) cm long, falcate or acidic rock types it is associated with Amentotaxus
nearly straight, 511 mm wide, subsessile to short yunnanensis, Cephalotaxus spp. and broad-leaved
petiolate and curved or asymmetric at base, tapering trees and shrubs (angiosperms) like Magnolia,
to an obtuse or acute-acuminate apex; margins flat or Quercus, and Rhododendron in montane evergreen
slightly revolute. Leaf texture coriaceous, with scler- or semi-deciduous forests.
eids causing a mottled, rugose upper surface, dark
green; on the lower surface two white bands bordered Uses
by wide green leaf margins and a midrib. Midrib
prominently raised on the adaxial (upper) side, lying The wood is used for tool making, furniture and
in a shallow groove, 0.81 mm wide, continuous to wood turning (handicrafts). This species is in cul-
the apex; on the abaxial side nearly flat and 1.21.5 mm tivation in China and was introduced to Europe
wide. Stomata on the abaxial surface in two bands from Hong Kong; it makes an attractive foliage
about as wide or slightly narrower than the green leaf shrub but only grows outside in warmer regions.
margins, numerous, randomly dispersed. Pollen cone This (and other) species is rare in gardens due to
racemes from large axillary or sub-terminal buds with poor availability in the horticultural trade. In China
25(10) together, 3.56.5 cm long, each with 1012 it is used as a bonsai plant. The seeds have high oil
cone pairs when full-grown; cones ovoid, 33.5 mm content and are surrounded by a very striking red
long; microsporophylls 68, peltate, each with 25 aril. Recent investigations have been undertaken to
pollen sacs. Seed-bearing structures near ends of analyse its potential for anti-cancer drugs similar to
foliage shoots, axillary, solitary on a slender, down- those found in Taxus.
curved, with 1.52 cm long peduncle, with 68(10)
decussate, keeled bracts enclosing a single, terminal 2 varieties are recognized:
ovule. Aril surrounding the seed ellipsoid or narrowly
obovoid, 2026 1013 mm, smooth, lustrous bright
red when ripe, with a mucronate apex. Seed proper Amentotaxus argotaenia (Hance) Pilg. var. argot-
much smaller, ca. 15 7 mm, oblong or ellipsoid, with aenia. Podocarpus argotaenia Hance, J. Bot. 21: 357.
a small mucronate apex. 1883; Nageia argotaenia (Hance) Kuntze, Revis.
Gen. Pl. 2: 800. 1891; Cephalotaxus argotaenia
Distribution (Hance) Pilg., in Engler, Pflanzenr. IV.5 [18]: 104.
1903. Type: China: Fujian, [in jugo Lo-fau shan,
China: Chongqing, Fujian, S Gansu, Guangdong, prov. Cantonensis], E. Faber s.n. [Acc. No. 22121],
Guangxi, Guizhou, Hong Kong, W Hubei, Hunan, Sep 1882, (holotype BM).
Jiangsu, NW Jiangxi, Central and SE Sichuan,
SE Xizang [Tibet], S Zheijiang; Taiwan; Indochina: Amentotaxus cathayensis H. L. Li, J. Arnold Arbor.
Lao PDR, N Viet Nam. 33: 195. 1952; Amentotaxus argotaenia (Hance) Pilg.
TDWG codes: 36 CHC-CQ CHC-GZ CHC-HU var. cathayensis (H. L. Li) P. C. Keng, [Chin. title; see
CHC-SC CHN-GS CHS-FJ CHS-GD CHS-GX CHS-HK Fl. Reipupl. Pop. Sin. 7: 452. 1978]: 2. 1957.
CHS-HN CHS-JS CHS-JX CHT 38 TAI 41 LAO VIE
Description Ecology
Leaves 311 cm long, 611 mm wide. Pollen cone This variety occurs on limestone karst mountains
racemes grouped with 24(5) together, rarely soli- on steep rocky slopes and ridges at around 1000 m
tary. Seeds including the aril 2026 mm long and altitude.
1013 mm wide.
Conservation
Distribution
Amentotaxus argotaenia var. brevifolia is known
China: Chongqing, Fujian, S Gansu, Guangdong, from a very limited area and has been severely
176 Guangxi, Guizhou, Hong Kong, W Hubei, Hunan, reduced in population size due to deforestation on
Jiangsu, NW Jiangxi, Central and SE Sichuan, the more accessible slopes.
SE Xizang [Tibet], S Zheijiang; Taiwan; Indochina: IUCN: CR (A2cd)
Lao PDR, N Viet Nam.
TDWG codes: 36 CHC-CQ CHC-GZ CHC-HU Uses
CHC-SC CHN-GS CHS-FJ CHS-GD CHS-GX CHS-HK
CHS-HN CHS-JS CHS-JX CHT 38 TAI 41 LAO VIE This variety grows to a small tree; its uses are similar
to those of the species but it is not known if it is in
Conservation cultivation in or outside China.
Vernacular names
Amentotaxus argotaenia (Hance) Pilg. var. brevi-
folia K. M. Lan & F. H. Zhang, Acta Phytotax. Assam catkin yew
Sin. 22 (6): 492. 1984. Type: China: Guizhou, Libo,
Dongjiang Shan, K. M. Lan & F. H. Zhang 82002 Description
(holotype GACP).
Trees to 20 m tall; trunk sometimes divided low
Description above the ground. Bark smooth, exfoliating in thin
flakes, whitish grey. Branches spreading or sweep-
Leaves 23.7 cm long, 57 mm wide. Pollen cone ing down. Foliage branchlets opposite, spreading
racemes grouped with up to 10 together, 1.55.5 cm at 2570 from the leading shoot axis, ascending
long. Seeds including the aril ca. 20 mm long and 10 or spreading, subterete or angular, with alternat-
mm wide. ingly twisted grooves running from one leaf base
to the next above, green in the first year, turning
Distribution from greenish yellow to yellowish brown in follow-
ing years, terminating in a conical bud with ovate-
China: SE Guizhou (Libo Xian). triangular, acute-mucronate scales. Leaves spreading
TDWG codes: 36 CHC-GZ distichously at 4580 from shoot axis, linear or
linear-lanceolate, (2)711(15) cm long, falcate or between 1600 m and 2000 m a.s.l. These are mossy
more or less S-shaped, (4)712 mm wide, short pet- forest dominated by Quercus spp., Castanopsis spp.,
iolate and curved or asymmetric at base, gradually Acer sp. and Rhododendron spp., with associated
tapering to an acute-acuminate apex, green with two taxa such as Magnolia sp., Michelia sp., Corylopsis
white or silvery grey bands bordered by wide green himalayana, Betula alnoides, Carpinus viminea and
leaf margins and a midrib on the lower surface; mar- Exbucklandia populnea. Amentotaxus assamica is a
gins flat or slightly revolute. Leaf texture coriaceous, rare species only known from two localities.
with a smooth or finely striated upper surface (lack-
ing sclereids in the leaf anatomy). Midrib promi- Conservation
nently raised on the adaxial (upper) side, lying in
a shallow groove, 0.40.7 mm wide; continuous to This species is known from two disjunct localities in 177
the apex, slightly raised and 11.8 mm wide on the Arunachal Pradesh, NE India, in only one of which
abaxial side. Stomata on the abaxial surface in two it has been collected recently (1980s). It is obviously
bands 1.52 times as wide as the green leaf margins, rare because several searches during the 1970s for it
numerous, randomly dispersed. Pollen cone racemes remained unsuccessful until it was found again in
from large axillary or sub-terminal buds with 24 1984. However, the region remains poorly explored
together, 45.5 cm long, each with 810 cone pairs botanically and we cannot say whether it is restricted
when full-grown; cones ovoid, 35 mm long; micro- to the localities known or more widespread. So far it
sporophylls 68, peltate, each with 24 pollen sacs. is not known from outside the state. Activities like
Seed-bearing structures axillary, solitary on 1.52.5 road building are opening up this border region to
cm long peduncles, with 8 decussate, ovate, keeled, settlement, leading to shifting agriculture and con-
2.55 mm long and 2.55 mm wide bract scales (the sequently forest disturbance and destruction. There
upper scales largest). Aril obovoid-oblong to ellip- is an urgent need to establish forest reserves as well
soid, 2535 1525 mm, lustrous green turning yel- as cultivation action to provide ex situ conservation
low to purple, with a mucronate apex. Seed proper for this species (Sahni, 1990).
much smaller, 1520 710 mm, oblong or ellipsoid, IUCN: EN [B1ab (iii) + 2ab (iii)]
with a small mucronate apex.
Uses
Taxonomic notes
No specific uses have been recorded of this species.
Ferguson (1985) has distinguished this species from
A. yunnanensis mainly on leaf anatomical charac-
ters. One of these, the lack of sclereids in the upper Amentotaxus formosana H. L. Li, J. Arnold Arbor.
mesophyll of the leaves, present in all other spe- 33: 196. 1952. Amentotaxus yunnanensis H. L. Li var.
cies, gives the leaves a smooth texture quite differ- formosana (H. L. Li) Silba, Phytologia 68: 25. 1990.
ent from its congeners. At the time, the seed bearing Type: Taiwan: Pingtung Co., [Taririku], S. Sasaki
structures were unknown, these were later described s.n. (holotype TAI). Fig. 38, 39
from newly collected specimens and appear to be
larger than in most other species, though variable. Etymology
Figure 1. Abies
alba in the Fort de
la Joux, Jura Mts.,
France
Figure 4. Abies
cephalonica flushing
leaves
Figure 5. Abies
delavayi var. delavayi
seed cones
Figure 7. Abies
delavayi var.
nukiangensis seed
cones
Figure 8. Abies
homolepis pollen cones
Figure 15. Abies pinsapo Figure 16. Abies procera seed cones
seed cone (photo W. Milliken)
Figure 18. Abies spectabilis Figure 19. Abies squamata bark of young
mature seed cone tree (photo P. de Spoelberch)
Figure 20. Abies squamata bark of old
tree (photo P. de Spoelberch)
Figure 22. Actinostrobus arenarius along Hwy. 123, Western Australia
Figure 33. Agathis lenticula tree at Figure 35. Agathis microstachya trees at Lake Barrine, Queensland
Kinabalu Park H.Q., Borneo
Figure 36. Agathis ovata trees near Yat, New Caledonia
Figure 34. Aga
this lenticula trunk
in Crocker Range,
Borneo
Figure 38. Amentotaxus
formosana tree in southern
Taiwan (photo C. N. Page)
Marywildea A. V. Bobrov & Melikyan, Komarovia 4: ding pollen. Microsporophylls imbricate, spreading
57. 2006. Type: Marywildea bidwillii (Hook.) A. V. at 4590 from a thin to stout rachis, apically free
Bobrov & Melikyan [Araucaria bidwillii Hook.]. at anthesis, divided into a stalk and a lamina; stalk
Titanodendron A. V. Bobrov & Melikyan, Komarovia 4: thin and weak, linear; lamina triangular, rhombic
60. 2006 Type: Titanodendron hunsteinii (K. Schum.) or peltate. Pollen sacs 620. Seed cones confined to 191
A. V. Bobrov & Melikyan [ Araucaria hunsteinii the mid and upper crown, axillary or terminal, ses-
K. Schum.]. sile or borne on a short, robust peduncle-like shoot,
subtended by modified leaves and then a transition
Name derived from the Araucanos (more correctly zone of sterile cone bracts, large, globose and heavy
known as Pehuenche) who live in the region of the (to 5, occasionally 10 kg), with milky resinous exu-
type species. date, usually ripening in second year. Bract-scale
complexes numerous, spirally arranged on a thick
Description rachis. Bracts broad, obtrullate, ovoid-oblong, cune-
ate or flabellate, often extended laterally into mem-
Small to very tall monoecious or more rarely dioe- branous (occasionally woody) wings, apical margin
cious trees with monopodial branching; branched thickened and with a narrow apophysis at tip. Seed
to ground when young but with clear bole when scale enclosed within and partly fused with bract,
old, resinous. Trunk terete, clothed with leaves when not or slightly wider than the seed, terminating in a
young. Crown columnar to candelabra- or umbrella- free scale-like ligule reaching more-or-less to base of
shaped. Bark exfoliating in plates and scales, or in the spur of the fertile bract. Ovules one (occasionally
horizontal strips. Branching conforming to either 2) per scale. Seeds cuneate; seed coat fused partly or
Massarts or Rauhs model depending on species wholly with scale, in one species (A. bidwillii) scale and
(sometimes intermediate between the two), often seed becoming separated; wing absent. Germination
reiterating. Apex of shoot a cluster of incompletely epigeal or hypogeal; cotyledons in 2 free and 2 fused
formed leaves; bud scales absent. Leaves persisting pairs, with 4 free, or 4 fused into 2 pairs at base.
for many years (in some species including on trunk).
All leaves similar or more often differentiated into 19 species in 4 sections.
more-or-less distinct juvenile and adult states. Leaves
spirally arranged on all axes including the trunk, Distribution
multi-veined, weakly to strongly keeled abaxially and
sometimes adaxially; apex incurved or not; margin South America: Argentina, Brazil, Chile, Paraguay;
entire or papillate or denticulate. Stomata amphisto- Australia: New South Wales, Norfolk Island,
matic (often very unequally, with fewer stomata abax- Queensland; New Caledonia; New Guinea.
ially). Resin canals in leaves alternating with vascular
bundles and in the same plane, or peripheral. Pollen Synopsis
cones solitary, terminal or axillary (lateral) on the
ultimate foliage branchlets, on different branchlets Recent phylogenetic analysis (Hollingsworth et al.,
to the seed cones and often lower down on the tree unpubl. data RBG Edinburgh) using various DNA
and/or on younger trees, subtended by a cluster of sequence data supports a taxonomic division of the
modified leaves, initially erect, frequently pendulous genus into 4 sections. These sections can also be rec-
when shedding pollen, cylindrical to ovoid, varying ognized by specific combinations of morphological
considerably in size between species and sometimes characters; some of which are given below with the
massive (up to 25 cm long), deciduous after shed- section names.
Sect. Araucaria. Dioecious or less commonly mon- well as intermediate shapes. Seedlings have differ-
oecious. Leaves undifferentiated into juvenile and ent leaves from saplings or young trees as well as
adult states, large, flat, thick or thin. Pollen cones mature trees. On mature trees adventitious branch-
axillary, often 2-several close together at tips of foli- ing often occurs and the leaves on these branches
age branches. Bract-scales nut-like, without wings, are quite similar, but not equal to those on young
indehiscent; seed retained on scale when shed. trees. The adult leaves described in the keys are those
Germination hypogeal. Cotyledons 2, long-stalked of the ultimate primary foliage branches of mature
in germination, stalks not fused. Seedling fleshy. trees that can produce cones and exclude those of
Species: A. angustifolia, A. araucana. adventitious branches. Under cultivation, trees may
retain (semi-)juvenile leaf shapes and sizes longer
192 Sect. Bunya Wilde & Eames. Monoecious. Leaves than in their natural habitat, especially when grown
undifferentiated into juvenile and adult states, large, under glass. It is virtually impossible to key these
flat, spreading or slightly imbricate. Pollen cones out, as juvenile leaves of many species are highly
axillary, often with 26 together. Seed cones axillary, similar. Pollen cones, as in the genus Agathis, have
subsessile or shortly pedunculate. Seeds only partly more informative characters than seed cones and are
fused with scale, becoming separated. Germination sometimes crucial for determination of species in
hypogeal. Cotyledons 2, remaining inside the seed addition (or in preference) to foliage. They can often
coat. Seedling fleshy. Species: A. bidwillii. been found under the tree (most species are monoe-
cious) and are then full size, having shed their pollen,
Sect. Eutacta Endl. Monoecious. Leaves clearly dif- and can be used for comparison of character states
ferentiated into juvenile and adult states; juvenile if not too far disintegrated. Trees grow a trunk and
leaves more-or-less narrow, falcate, spreading; adult branches in specific ways that may be distinct among
leaves scale-like and imbricate, adpressed. Pollen taxa. The architecture of branching has been studied
cones terminal. Seed cones terminal, long-pedun- and models are named after their describers. Two
culate. Bract-scales samara-like, thinly winged, of these models apply to the species of Araucaria,
indehiscent; seed retained on scale when shed. Massarts and Rauhs. In both, there is a single, domi-
Germination epigeal. Cotyledons 4, completely free nant, erect leader making the stem (trunk in mature
or fused into two pairs at base, freed from seed at trees) and branching occurs in pseudo-whorls at
germination. Seedling not fleshy. Species: A. berni- regular intervals on it. The branches spread out more
eri, A. biramulata, A. columnaris, A. cunninghamii, or less horizontally. In Massarts model they remain
A. heterophylla, A. humboldtensis, A. laubenfelsii, A. horizontal, as do the lateral branches, or they may be
luxurians, A. montana, A. muelleri, A. nemorosa, A. more or less drooping, and buds facing up and down
rulei, A. schmidii, A. scopulorum, A. subulata. on each subsequent pseudo-whorl remain dormant.
In Rauhs model, the branches are curved upward
Sect. Intermedia C. T. White. Monoecious. Leaves towards their ends and all buds on a pseudo-whorl
differentiated into juvenile and adult states, juve- can grow to form new branches, repeating the pat-
nile leaves small, needle-like, flat; adult leaves large, tern on the stem. Adventitious branching, often trig-
flat, spreading, sometimes slightly imbricate. Pollen gered by some damage, usually does not conform to
cones axillary. Seed cones axillary. Bract-scales the model and can disrupt the architecture, so only
samara-like, with broad relatively thin wing, indehis- primary branching should be considered.
cent; seed retained on scale when shed. Germination
epigeal. Cotyledons 2, free, freed from seed at ger- 1a. Leaves differentiated into juvenile and adult
mination. Seedling not fleshy. Species: A. hunsteinii. stages. Bract scales of seed cones samara-like.
Seedlings not fleshy 2
Key to the sections of Araucaria 1b. Leaves undifferentiated into juvenile and adult
stages. Bract scales of seed cones nut-like or
The leaves of many species in the genus Araucaria large and heavy. Seedlings fleshy 3
are highly variable and part of this variation is asso- 2a. Adult leaves imbricate, often adpressed. Coty
ciated with the age of the branch on which they ledons of seedlings 4, free or sometimes fused
appear. There are often juvenile and adult forms, as into two pairs at base Sect. Eutacta
2b. Adult leaves spreading. Cotyledons of seed- 4a. Adult leaves nearly triangular. Crown colum-
lings 2, free nar or broadly conical. Mature seed cones
Sect. Intermedia; a single species, 1013 79 cm; bract scales 2225 1520 mm
A. hunsteinii A. laubenfelsii
3a. Bract scales of seed cones nut-like, without 4b. Adult leaves broadly ovate. Crown domed, very
wings, indehiscent; seed retained on scale when open. Mature seed cones 811 79 cm; bract
shed Sect. Araucaria scales 2535 1825 mm A. montana
3b. Bract scales of seed cones large, heavy, with 5a. Branching architecture according to Rauhs
woody wings, dehiscent; seed shed from scale model, without or with infrequent adventitious
at maturity branching, developing an open crown in
Sect. Bunya; a single species, A. bidwillii mature trees A. biramulata 193
5b. Branching architecture according to Massarts
Key to the species of Section Araucaria model, often with frequent adventitious branch-
ing, developing a more or less dense crown in
1a. Foliage branches with and including adult mature trees 6
leaves up to 40 mm wide. Pollen cones 1530 6a. Adult leaves ovate or broadly ovate with a
mm wide. Resin exuding from bark on trunk rounded apex, 416 2.56.5(8) mm. Pollen
reddish A. angustifolia cones (10)1215(17) cm long and 2028 mm
1b. Foliage branches with and including adult wide A. luxurians
leaves 6090 mm wide. Pollen cones 4050 6b. Adult leaves from subulate-acicular to broadly
mm wide. Resin exuding from bark on trunk ovate with an acute, acuminate or obtuse, never
clear, drying white A. araucana rounded apex, 210 1.25(6) mm. Pollen
cones 212 cm long and 322 mm wide 7
Key to the species of Section Eutacta 7a. Adult leaves subulate or lanceolate 8
7b. Adult leaves (ob)ovate or elliptic-ovate 11
1a. Foliage branches with and including adult 8a. Pollen cones very slender, 37 mm wide; micro-
leaves (12)1550 mm wide; branching archi- sporophylls connate. Outer bark on trunks
tecture according to Rauhs model 2 exfoliating with thin, curling flakes
1b. Foliage branches with and including adult A. cunninghamii
leaves (2.5)313(16) mm wide; branching 8b. Pollen cones 720 mm wide; microsporophylls
architecture mostly according to Massarts imbricate or divergent. Outer bark on trunks
model (in one species Rauhs model) 5 exfoliating in horizontal strips 9
2a. Foliage branches with and including adult 9a. Pollen cones 25 cm long, 711 mm wide. Seed
leaves 3050 mm wide; adult leaves 2032 cones when mature 79 56 cm. Foliage
9.518.5 mm, triangular to linear-triangular branches with and including adult leaves 610
A. muelleri mm wide. Endemic to the summit of Mt. Pani
2b. Foliage branches with and including adult in New Caledonia A. schmidii
leaves (12)1533 mm wide; adult leaves 820 9b. Pollen cones 512 cm long, 1020 mm wide.
(25) 414 mm, broadly lanceolate, ovate or Seed cones when mature 7.512 610 cm.
nearly triangular 3 Foliage branches with and including adult
3a. Trees with sparse but long branches; crown leaves 4.58(12) mm wide. Occurring else-
candelabra shaped, with tufts of thick foliage where in New Caledonia 10
branches at ends; adult leaves broadly lanceo- 10a. Microsporophylls of pollen cones divergent,
late, short retained on leading branches spreading at nearly 90 from the rachis, ovate,
A. rulei 58 45 mm. Adult leaves 410 mm long
3b. Trees with numerous branches; crown colum- A. nemorosa
nar, broadly conical or domed; foliage branches 10b. Microsporophylls of pollen cones imbricate,
usually not limited to ends of main branches; spreading at 45 from the rachis, triangular, 34
adult leaves broadly ovate to nearly triangular, 22.5 mm. Adult leaves (3)46 mm long
long retained on leading branches 4 A. subulata
11a. Crown of mature trees domed to flat-topped. Vernacular names
Microsporophylls of pollen cones connate, 2.5
2 mm A. scopulorum Parana pine, Candelabra tree; pinheiro-do-Paran,
11b. Crown of mature trees columnar or narrowly also recorded are: pinhao, pinheiro, pinho-nacional
conical. Microsporophylls of pollen cones (Portuguese); pino Paran (Spanish); curiy or kuriy
imbricate, 37(10) 24 mm 12 (perhaps originally Amerindian) and many other
12a. Foliage branches with and including adult variants denoting ecotypes or seed types.
leaves very slender, 37 mm wide; adult leaves
broadly ovate, 2.54.7 1.52.5 mm Description
A. bernieri
194 12b. Foliage branches with and including adult Dioecious or less commonly monoecious trees to
leaves slender to more robust, (3)512 mm 50 m tall, to 2 m d.b.h.; trunk straight. Bark to 15
wide; adult leaves elliptic-ovate or ovate, 48 cm thick, rough and deeply fissured, exfoliating in
1.85(6) mm 13 small chips and plates, sometimes exposing smooth
13a. Crown of mature trees columnar but open, in patches of reddish inner bark; outer bark grey. Resin
older trees with a flat or candelabra shaped top, exudate reddish. Crown in mature trees domed and
in young trees pyramidal. Pollen cones small, finally flat-topped, with branches only at the top
33.5 cm long and 910 mm wide of the tree; branching according to Rauhs model.
A. humboldtensis Primary branches in pseudo-whorls of 48, up to 5
13b. Crown of mature trees narrowly columnar or m long, spreading or assurgent, mostly with second
narrowly conical, with a narrowing top, in to third order branches, caducous. Adventitious foli-
young trees (narrowly) conical. Pollen cones age branches uncommon, usually associated with
(2.5)47(10) cm long and 1022 mm wide damage. Foliage branchlets spreading or ascend-
14 ing (sometimes more or less pendent) on primary
14a. Crown of mature trees narrowly columnar. branches, forming large tufts of foliage, of unequal
Pollen cones 1322 mm wide; microsporophylls length up to 50 cm, up to 40 mm wide (including
oblong-triangular, 57(10) 2.53(4) mm leaves), but of unequal width from base to apex
A. columnaris depending on size and spread of leaves, more or
14b. Crown of mature trees narrowly conical. Pollen less flexible. Adult leaves imbricate or variously
cones 1015 mm wide; microsporophylls rhom- spreading (4090), decurrent, ovate to lanceo-
bic, 34 4 mm A. heterophylla late, 1.55 cm 320 mm, varying much in length
and width often on a single branch, flat or slightly
concave adaxially, sometimes weakly keeled abaxi-
Araucaria angustifolia (Bertol.) Kuntze, Rev. Gen. ally, smooth or striate, tapering to a pungent apex;
Pl. 3: 375. 1898. Columbea angustifolia Bertol., margins entire. Stomata in 4050 indistinct rows
Opusc. Sci. 3: 411. 1819. Type: Brazil: Rio de Janeiro, on the abaxial surface and slightly fewer but more
Corcovado, [In Brasilia legit G. Raddi], G. Raddi conspicuous rows on the adaxial surface. Pollen
s.n. (holotype BOLO). Pl. 6 cones axillary at end of foliage branchlets, solitary
or in small clusters, initially erect, cylindrical and
Araucaria angustifolia (Bertol.) Kuntze var. depen- green, becoming pendulous, elongated, curved and
dens J. R. de Mattos, Loefgrenia 21: [2]. 1967. pinkish brown at anthesis, 815 cm 1530 mm.
Araucaria angustifolia (Bertol.) Kuntze var. vinacea Microsporophylls connate, spreading at ca. 90 from
J. R. de Mattos, Loefgrenia 111: 1. 1997. a stout rachis, apically free at anthesis; stalk thin and
weak, linear, 35 mm long; lamina cuneate, rhombic
Etymology in outline, 45 3 mm, lateral margins entire; apex
curved; abaxial surface smooth. Pollen sacs 812(
The species epithet alludes to the (often) narrower 15), linear, 68 mm long. Seed cones axillary on stout
leaves than those of A. araucana, the only other spe- foliage branches which widen below the cone, usu-
cies native to South America. ally 35 together in whorls, erect. Immature cones
195
plate 6. Araucaria angustifolia. 1. Habit of tree. 2. Foliage branch with pollen cone. 3. Leaf. 4. Pollen
cone. 5. Microsporophyll with pollen sacs. 6. Seed cones, one with top removed to show seeds.
ovoid, 46 cm long, densely covered with recurved The climate varies with latitude and altitude; on
bract tips, green, maturing in 2022 months. Mature the Planaltos at around 1000 m frost is a common
cones chestnut brown, ovoid-globose to globose, occurrence in winter with minima to 10 C while
usually slightly longer than wide, (8)1220 818 snowfall, although rare, can occur in most of the
cm (green weight 34 kg). Bracts ovoid-oblong to range of the species. According to Golte (1993) the
cuneate, more or less angular, ca. 5 2 cm includ- northern limit of natural occurrence of A. angustifo-
ing vestigial wings if present, distally thickened to a lia coincides with the zone where tropical winter dry
more or less rhombic, transversely keeled apophy- seasons begin to have influence upon the vegetation.
sis, ending in a caudate, slightly curved, 810 mm Rains are mostly summer rains in much of its range,
long tip. Seed scale not wider than the seed; ligule but with more winter rains in the south, to a total of
196 fragile, 11.5 mm long, sometimes absent. Seeds 14002500 mm per annum. The prevalent soil type
ovoid-oblong, ca. 4 1.5 cm, not flattened, striated in the Araucaria forests is a well-drained, iron-rich
longitudinally or smooth, ripening lustrous red- red-yellow podzol with a low pH value.
brown (sometimes striped darker red, or with white
or pale apex, or occasionally whole seed very pale). Conservation
Papua New Guinea. In the central mountain range Araucaria hunsteinii has been grown as a plantation
from an isolated population on the Wamira River in crop in New Guinea since 1948. The wood is used
the east through the Owen Stanley Range and the in the local sawmilling and plywood industries and
Bismarck Range, with other isolated stands near for making aircraft frames. Fires destroyed much
Sattelburg in the Huon Peninsula and on the Tagari of the Bulolo plantation in 1997. Klinki pine plan-
River in the Central Highlands. tations have also been established on tin tailings in
TDWG codes: 43 NWG-PN Malaysia and more recently in Costa Rica. The lat-
ter plantations were set up as part of a controversial
210 Ecology programme, KLINKIFIX, which aims to mitigate
greenhouse gas emissions.
This species forms groves or stands, or occurs scat-
tered in tropical montane monsoon forests on moist
sites usually in inter-montane valleys. Its altitudi- Araucaria laubenfelsii Corbasson, Adansonia, sr. 2,
nal range is (550)7501700(2100) m a.s.l. Mature 8: 467. 1969. Type: New Caledonia: Grande Terre,
trees are canopy emergents above angiosperm trees, Province Sud, on ridge between Mts. Dzumac and
effectively forming an upper canopy layer. This Mt. Ouin, H. S. MacKee 16441 (holotype P). Fig. 51
species occurs in two types of forest, a drier and a
wetter one; in the drier type the canopy of angio- Etymology
sperms reaches only 1525 m of average height, with
A. hunsteinii attaining twice that height. Associated This species was named after David J. de Laubenfels,
common evergreen tree species in this forest type specialist on gymnosperms especially the Podocar
are Aleurites moluccana, Celtis sp., Heritiera sp., paceae and the Araucariaceae.
Macaranga sp., and Pouteria luzonensis; deciduous
species are Garuga floribunda, Protium macgrego- Vernacular names
rii, Sterculia sp., and Terminalia sp. In high rain-
fall localities both angiosperms and A. hunsteinii ouaou (name recorded on H. S. MacKee 30370, P).
become much taller, with the conifer towering to
6090 m. Common associated canopy tree spe- Description
cies are Acmena sp., Elmerillia papuana, Flindersia
amboinensis, F. pimenteliana, Pometia pinnata, and Monoecious trees to 30 m tall, to 1 m d.b.h.; trunk
Xanthophyllum papuanum. Cerbera floribunda, straight. Bark to 4 cm thick, exfoliating in horizon-
Cryptocarya sp., Dysoxylum sp., Gnetum gnemon, tal strips; inner bark reddish to pink; outer bark
Litsea sp., and Myristica sp. are common in the sub- brown or grey. Resin exudate clear or yellow, dry-
canopy tree layer. Soils are neutral to acidic with a ing white or grey. Crown in mature trees colum-
high clay content. Precipitation ranges from 800 nar or broadly conical, open, branched along the
mm to more than 4000 mm per year. greater part of the trunk; branching according to
Rauhs model. Primary branches followed by adven-
Conservation titious branching lower on the trunk. Primary first
order branches of mature trees in pseudo-whorls of
This species was graded LR(nt) by Farjon & Page 45, 12.5 m long, spreading nearly horizontally to
(1999), which was still the rating (NT) in the 2008 assurgent, caducous. Foliage branchlets assurgent
IUCN Red List of Threatened Species (www.iuc- or nearly erect on primary branches in mature trees
nredlist.org) although the destruction of so many and confined to distal parts, of more or less equal
trees in recent forest fires may mean that the species length (3545 cm) on either side of the axis branch
needs to be reassessed. The recalcitrant seeds cannot and shortening towards the end, (12)2033 mm
be stored for long periods making propagation from wide (including leaves). On mature trees leaves of
seed difficult. two kinds: adult leaves and semi-juvenile leaves
IUCN: NT on adventitious branches. Adult leaves on ultimate
branchlets nearly triangular, 820(24) 412 mm, dense forest, in which the surviving araucarias
at least twice as long as wide, similar in size along become large and tall emergents. It appears therefore
length of branchlet, strongly convex, usually with that this species is dependent on episodic fires for its
5 longitudinal ridges alternating with grooves or perpetuation in the vegetation.
sometimes smooth; margins incurved; apex acute
or obtuse. Stomata abaxially in bands located in the Conservation
grooves between the ridges; adaxially in numerous
continuous rows from base to apex. Semi-juvenile The threats are those common to most of the New
leaves irregularly spreading at 4090, acicular to Caledonian species: fires, land management of non-
lanceolate, 1325 48 mm, tapering to an incurved, agricultural areas, and clear-cutting. Most popula-
acute apex. Pollen cones terminal, solitary, initially tions occur on serpentine and some, especially in 211
erect but becoming pendulous and curved at anthe- the Province Nord, are subject to nickel mining. The
sis, 811(15) 23 cm. Microsporophylls imbricate, species has probably now disappeared from one of
spreading at 45 from a slender rachis; stalk thin and such sites, Mt. Kaala (pers. observation, Nov 2005).
weak, 35 mm long; lamina nearly triangular, 46 A reassessment of its status seems necessary.
35 mm, thick and firm; lateral margins entire; apex IUCN: NT
obtuse; abaxial surface smooth. Pollen sacs 1012,
thin, straight, 56 mm long. Seed cones terminal Uses
on short, stout foliage branches, usually solitary,
sometimes 23 together, erect, broadly ovoid to sub- The wood of this species is not exploited and no
globose, 1013 79 cm. Bracts flabellate, 2.22.5 other uses are recorded.
1.52 cm including rounded, thin membranous
wings, distally thickened and transversely keeled,
ending in a rostrate, upturned, 1015 mm long, acute Araucaria luxurians (Brongn. & Gris) de Laub.,
tip. Seed scales not wider than the seed; ligule frag- Fl. Nouv. Caldonie Dpend. 4: 92. 1972. Araucaria
ile, 1.52 mm long. Seeds cuneate, 1416 68 mm, cookii R. Br. ex Lindl. var. luxurians Brongn. & Gris,
smooth, yellowish brown when ripe. Ann. Sci. Nat. Bot., sr. 5, 13: 354. 1871; Araucaria
columnaris (G. Forst.) Hook. f. luxurians (Brongn.
Distribution & Gris) E. H. Wilson, J. Arnold Arbor. 7: 84. 1926.
Type: New Caledonia: Grande Terre, Province Sud,
New Caledonia: Grande Terre. In Province Nord Mt. Canala, B. Balansa 2510 (lectotype P).
only known from le Art and Mt. Kaala; more abun-
dant in Province Sud. Etymology
TDWG codes: 60 NWC
The species epithet (Latin luxurians = luxuriant)
Ecology refers to the branching habit.
New Caledonia: Grande Terre, Province Nord, sum- Araucaria bernieri J. T. Buchholz var. pumilio Silba, J.
mit of Mt. Pani. Int. Conifer Preserv. Soc. 7 (1): 21. 2000.
TDWG codes: 60 NWC
Etymology
Ecology
218 The species epithet comes from the Latin scopulo-
Araucaria schmidii grows in dense humid montane rum = of cliffs or crags.
cloud forest on steep slopes and exposed ridges at the
edge of the summit plateau of Mt. Pani, on micas- Vernacular names
chist substrate and often directly from rock crevices.
Its altitudinal range is 14001630 m a.s.l. This is the No common names have been recorded for this
only New Caledonian Araucaria that grows exclu- species.
sively on non-ultrabasic substrates. At its lowest
limit on the mountain this species is still accompa- Description
nied by Agathis montana, whereas near the summit
it is the only tall tree, which is emergent above a Monoecious trees to 20 m tall, to 0.5 m d.b.h.; trunk
dense undergrowth of evergreen angiosperm shrubs straight. Bark to 2 cm thick, exfoliating in horizon-
and small trees, as well as tree ferns (Cyathea vie- tal scales; inner bark reddish; outer bark grey with
illardii) and small palms. Mt. Panis eastern slopes nearly white patches. Resin exudate clear or yel-
receive the highest rainfall on the island, estimated low, drying white or grey. Crown in mature trees
to exceed 8000 mm per year (Schmid, 1989; cited in irregular, open, branched along greater part of
Golte, 1993). trunk, domed to flat-topped in older trees; branch-
ing according to Massarts model. Primary first
Conservation order branches of mature trees in pseudo-whorls
of 57, 12.5 m long, spreading nearly horizon-
The only confirmed population of this species, a tally to ascending, caducous. Adventitious foliage
few hundred trees, is that at Mt. Pani. It is the most branches similar to primary branches on mature
inaccessible of all species due to its location on the trees but shorter. Foliage branchlets ascending or
summit of the highest mountain in New Caledonia, nearly erect near ends of primary branches, form-
to which a primitive, very steep footpath through ing two pectinate rows, 1020 cm long and short-
dense rain and cloud forest gives the only access. ening towards the end, 49 mm wide (including
There is no prospect of mining due to the geology of leaves), changing thickness along a single branch-
this mountain massif. It occurs within an established let due to variable leaf sizes, flexible but standing
nature reserve, for which there are expansion plans nearly erect. On mature trees leaves of two kinds:
and, in future, to include it with the coastal reefs adult leaves on primary branches and semi-juvenile
in a UN World Heritage Site (Henry Blaffart, pers. leaves on adventitious branches. Adult leaves on
comm., 2005). ultimate branchlets ovate or obovate, 2.55.5(6)
IUCN: VU (D2) 2.53.5 mm, keeled abaxially; apex incurved, sub-
acute; leaves subtending pollen cones spreading,
Uses straight, narrowly oblong-lanceolate, narrower than
lower leaves on same branchlet and 23 times lon-
The wood is not exploited and no other uses have ger. Stomata mainly in 12(3) rows nearest the keel
been recorded for this species. on each abaxial face, extending to apex, on the
adaxial face in ca. 20 rows from near base to apex. Conservation
Semi-juvenile leaves narrowly obovate to elliptic,
3.58 1.22.3 mm, curved and with an abaxial keel. The distribution of this species nearly coincides with
Pollen cones terminal, solitary, initially straight but nickel mining areas and many populations are in or
curved at anthesis, cylindrical, 2.55.5 cm 1012 adjacent to active nickel mines. When compared
mm. Microsporophylls spreading at 45 from a stout with the distribution map given by the Laubenfels
rachis; stalk thin and weak, 23 mm long; lamina (1972) recent distribution data seem to register an
ovate-triangular, appearing rhombic when con- increase, but this is an artefact due to more complete
nate, 2.5 2 mm; apex acute. Pollen sacs 68, linear, data, including collections made since 1972. In real-
straight, 3 mm long. Seed cones terminal on short, ity, numbers of trees and probably numbers of (sub-)
stout foliage branches which widen below the cone, populations are declining. Fire and erosion have also 219
usually solitary, erect, broadly ovoid to subglobose, been noted as threats. None of the known popula-
5.59 4.57.5 cm. Bracts flabellate, 23(3.4) 1.8 tions of this species are within a protected area at
2.5 cm including rounded, thin membranous wings, present.
distally thickened and transversely keeled, ending IUCN: EN [B1ab(iiv)+2ab(iiv)]
in a 46 mm long, upturned, acute tip. Seed scales
not or slightly wider than the seed; ligule fragile, Uses
1.52 mm long. Seeds cuneate, 1720(24) 69
(10) mm, smooth, light brown when ripe. The wood is not exploited and no other uses are
recorded in New Caledonia. This species is in culti-
Taxonomic notes vation in Tasmania.
plate 7. Araucaria scopulorum. 1. Habit of tree. 2. Foliage branch with pollen cones. 3. Section of branchlet
with leaves. 4. Pollen cone. 5. Microsporophyll with pollen sacs. 6. Seed cone. 7. Seed cone scales.
branches of mature trees in pseudo-whorls of 57(10) Ecology
mostly restricted to the top of the tree, 0.51.5 m
long, spreading or ascending, soon caducous. Araucaria subulata is emergent in dense humid sub-
Adventitious foliage branches numerous. Foliage montane to montane rainforest, at altitudes between
branchlets assurgent near ends of primary branches, 150 m and 1070 m a.s.l. It grows sometimes on cliffs;
of more or less equal length (3050 cm) in two pecti- apparently always on ultramafic soils or rubble
nate rows and shortening towards the end, 4.58 mm derived from peridotite or serpentine. It occurs fre-
wide (including leaves), remaining of equal width quently in ravines and steep valleys on the rainward
from base to apex, flexible and whip-like. On mature side (E and SE) of mountains, with accumulation of
trees leaves of two kinds: adult leaves on primary litter and humus. This species is often co-dominant
branches and semi-juvenile leaves on adventitious with Agathis lanceolata and Montrouziera cauliflora 221
branches. Adult leaves on ultimate branchlets subu- (Guttiferae) and emerges from a 1230 m tall canopy
late to broadly lanceolate, (3)46 12.5(2.7) mm, of numerous evergreen angiosperms, among which
very glossy abaxially, strongly incurved, acute, acu- sometimes Nothofagus sp. is abundant. Podocarpus
minate or obtuse. Stomata on the abaxial surface sylvestris is another frequent coniferous tree in this
near leaf base, on the adaxial surface in several rows forest type. Tree ferns (Cyathea sp.) and palms are
for most of leaf length. Semi-juvenile leaves awl- also common, the former especially in clearings.
shaped or subulate, 48.5 0.51.2 mm; apex obtuse,
non-pungent. Pollen cones terminal, cylindrical, Conservation
initially erect and straight, pendulous and curved
at anthesis, 510 cm 1214 mm. Microsporophylls This species is relatively common in the south,
imbricate, spreading at 45 from a stout rachis; stalk where it is associated with (remnants of) sub-mon-
thin and weak, 5 mm long; lamina more or less tri- tane rainforest. Outside protected areas, however,
angular, 34 22.5 mm; apex acute or acuminate. this type of forest is often threatened by deforesta-
Pollen sacs ca. 10, linear, 2.53 mm long. Seed cones tion and fires from already deforested surrounding
terminal on short, stout foliage branches which areas. Mining is potentially of concern although at
widen below the cone, solitary, erect, subglobose or present no mining operations are known to exist in
broadly ovoid, 7.512 6.510 cm. Bracts flabellate, any of its localities. Several populations are within
2.53 2.22.7 cm including rounded, thin membra- protected areas.
nous wings, distally thickened but thin-edged, end- IUCN: NT
ing in a caudate, upward curved, 810 mm long tip.
Seed scales not wider than the seed; ligule fragile, Uses
11.5 mm long. Seeds cuneate-oblong, 1720 89
mm, striated longitudinally or smooth, light brown The wood is of high quality and has been locally
when ripe. exploited for use in carpentry.
Distribution
Greek: athroos = crowded; taxis = arrangement; 2a. Leaves 412 23 mm, mostly connate but with
referring to the leaves or seed cone scales. free apex A. laxifolia
2b. Leaves 718 34 mm, the distal part spreading
Description A. selaginoides
Distribution
Athrotaxis laxifolia Hook., Icon. Pl., n.s., 2: t. 573.
Australia: Tasmania, mainly on the Central Plateau, 1843. Type: Australia: Tasmania, R. C. Gunn [369]
the Great Western Tier and westward mountains, s.n. (holotype K).
more scattered in the S of the island.
TDWG codes: 50 TAS Etymology
Latin: australis = southern; cedrus = cedar; therefore directed forward, covered by leaves, persistent, turn-
southern cedar. ing orange- to red-brown, soon grey. Leaves scale-
like, decussate, in opposite ranks, dimorphic; facial
Description leaves very small, ca. 1 mm, partly hidden by larger
laterals, incurved-appressed, keeled, acuminate or
See the species description. obtuse; lateral leaves decurrent, bilaterally flattened,
226 34.5 mm long on ultimate branchlets, incurved;
Distribution apex free or appressed, obtuse; margins entire; leaves
(nearly) hypostomatic; stomata in depressed green-
As for the species. ish white bands bordered by thick green margins on
the abaxial side and in 2 small groups near base on
the adaxial side; transitional leaf form on leading
Austrocedrus chilensis (D. Don) Pic. Serm. & shoots (whip shoots) long decurrent (to 15 mm); lat-
Bizzarri, Webbia 32 (2): 482. 1978. Thuja chilensis erals with a spreading, curved distal part terminat-
D. Don, in Lambert, Descr. Pinus, ed. 8, 2: 128. ing in an acute apex. Pollen cones terminal, solitary,
1832; Libocedrus chilensis (D. Don) Endl., Syn. 57 23 mm; microsporophylls 1220, decussately
Conif.: 44. 1847. Type: Chile: [Habitat in Andium arranged, imbricate, subpeltate with acuminate
Regni Chilensis montibus], J. A. Pavn y Jimnez apex; pollen spherical, in 34 abaxial pollen sacs
[ex herb. A. B. Lambert] s.n. (lectotype K). near the lower margin of the microsporophyll. Seed
Fig. 58, 59 cones terminal, often numerous on ultimate branch-
lets with non-modified leaves, maturing in one year
Etymology to ovoid-oblong, apically slightly flattened cones
714 47 mm. Bract-scale complexes 4, decussate,
The species epithet refers to its country of origin. in pairs of very unequal size, the upper scales fertile
and twice as large as the lower, ovoid-oblong, with
Vernacular names a subapical curved umbo (bract tip), opening late,
becoming woody, longitudinally striated or grooved
Chilean cedar, Cordilleran cypress; ciprs de la abaxially, smooth adaxially, brown. Columella coni-
Cordillera, Ciprs (Spanish) cal to more or less clavate, 23 mm long. Seeds 24
per cone, angular-ovoid, ca. 4 2.5 mm, brown with
Description light yellowish hilum and 2 wings, one rudimentary,
the other well developed, angular-oblong, 68
Shrubs or trees to 1520(25) m tall, evergreen, dioe- 34 mm.
cious, rarely monoecious; trunk multistemmed or
monopodial to 1.5 m d.b.h. Bark smooth, soon flak- Taxonomic notes
ing, orange-brown turning grey with brown inner
bark, fissued and scaly, exfoliating in long strips. First described as a species of Thuja, this taxon was
Branches numerous, spreading, assurgent in higher transferred to Libocedrus by Endlicher (1847), then
part of crown, persistent, forming a conical or pyra- to a monospecific genus Austrocedrus by Florin &
midal or more rounded crown in trees. Foliage Boutelje (1954), who in creating the genus failed to
branches dense, spreading in horizontal planes cite the correct basionym. This error was corrected
or assurgent, nearly erect in young trees, ultimate by Pichi Sermolli & Bizzarri, who thereby validated
branchlets plagiotropic, opposite or nearly so, short, the currently used combination as cited above.
Distribution Patagonian steppe it is the only tree species, but does
not attain large size there.
Central and S Chile: Biobio, La Araucania, Los
Lagos, Maule, OHiggins, Santiago; S Argentina: Conservation
Chubut, Neuqun, Rio Negro.
TDWG codes: 85 AGS-CB AGS-NE AGS-RN CLC-BI This species, though still wide spread, is under vari-
CLC-LA CLC-MA CLC-OH CLC-SA CLS-LL ous pressures that make it potentially vulnerable to
extinction. Felling, expansion of human habitation
Ecology and agriculture, transformation of natural forest
to plantation forest with exotic trees (Eucalyptus,
This species has a natural distribution accross Pinus), and fires have all played a role in the decline 227
nearly 12 degrees of latitude in the Andes, from 32 of its area of occupation. A dieback problem has
to 4244 S, thus ranging from a Mediterranean been reported in the Argentinian populations
type climate to the cold montane Patagonian steppe (Havrylenko et al., 1989). Seed viability is often
climate. It generally shifts from N to S from the low (down to less than 5%) due to insect predation
western slopes to the eastern slopes of the Andes, (M. Gardner, pers. comm., 1996). Only a relatively
leaving more mesic conditions to other tree spe- small proportion of its populations is within pro-
cies. It often occurs in pure stands or mixed with tected areas in both countries (ca. 15% in Argentina).
Nothofagus dombeyi after quickly covering former Grazing of livestock and feral deer is a problem both
disturbance areas (fire, earthquakes, volcanic depos- outside and inside these reserves. Large areas in
its) but also grows associated with Fitzroya cupres- which this species is abundant appear to exist in the
soides in the coastal ranges and with Nothofagus Lake District of Argentina (AF pers. obs. Dec 2012).
spp. in the Andes. Austrocedrus chilensis gradually IUCN: NT
disappears in the wetter regions to the south where
Nothofagus will eventually dominate after distur- Uses
bances. Its tolerance to shade is relatively low, but
apparently somewhat higher than of its common Timber of Chilean cypress is used for general car-
associate Nothofagus dombeyi, so that some individ- pentry and furniture. Its fragrant and durable quality
uals can persist in the understorey. It is longer-lived makes it especially suitable for panelling and for out-
than its competitor, though not reaching more than door furniture. It is a useful species for horticulture,
1000 years, usually around 500 years, so it can fill as in cultivation it usually grows a fastigiate, columnar
gaps left by fallen Nothofagus dombeyi. Its edaphic habit (an as yet unexplained phenomenon seen also
requirements are lower than for most tree species in in other members of Cupressaceae, e.g. Calocedrus
the region, so it can occupy sandstone, volcanic, or decurrens). Despite this it is still uncommon and
granitic outcrops where other species will not estab- largely confined to arboreta and other botanical col-
lish, thereby extending its range into the wetter lati- lections. A few cultivars have been mentioned in the
tudes of the Southern Andes. Where it borders the 19th century but may no longer be available.
Austrotaxus R. H. Compton, J. Linn. Soc., Bot. 45: 427. 1922. Type: Austrotaxus
spicata R. H. Compton (Taxaceae).
Latin: australis = southern; taxus = yew; therefore ing to an acute apex; coriaceous, lustrous dark green
southern yew. above, light green below; margins slightly revolute.
Midrib forming a narrow groove on the adaxial
Description (upper) face from base to apex, on the abaxial side
raised, 0.81.2 mm wide. Stomata in two broad bands
See the species description. of numerous wavy, intermittent lines on tha abaxial
228 side only, bordered by narrow green margins and the
Distribution midrib. Pollen cones axillary, solitary but aggregated
near base of new foliage shoots, on a scaly short
As for the species. peduncle or dwarf shoot, elongating to 1215 mm,
catkin-like, creamy white, microsporophylls1218,
remote, spirally arranged, reniform, concave, on the
Austrotaxus spicata R. H. Compton, J. Linn. Soc., adaxial side bearing two or three rows of globose
Bot. 45: 427. 1922. Type: New Caledonia: Grande pollen sacs, up to 8 sacs in the outer row and 23
Terre, Province Nord, Mt. Ignambi, R. H. Compton larger ones in the inner row. Seed-bearing structures
1155 (holotype BM). Fig. 60 axillary, solitary near base of new foliage shoots, on a
very short, scaly peduncle or dwarf shoot, the upper
Etymology scales enlarged, imbricate, triangular, enclosing a
single apical ovule. Aril almost entirely enclosing the
The species epithet (Latin spicatus = spike) refers to seed, ovoid and slightly flattened when mature, 1215
the pollen cones. mm long, 79 mm wide, smooth, glaucous green
ripening to orange, only leaving the seed apex free.
Vernacular names Seed proper ovoid-oblong, yellowish brown with a
darker apex and minute micropylar tip.
New Caledonia yew.
Distribution
Description
New Caledonia: Grande Terre.
Trees to 25 m tall; trunk erect and straight to 60 cm TDWG codes: 60 NWC
d.b.h. in forest. Bark fibrous, exfoliating in short
strips or on the lower trunk becoming tesselated, Ecology
dark reddish brown weathering grey. Branches
numerous, forming a dense crown. Foliage branch- Austrotaxus spicata is locally common in the mon-
lets subopposite or more or less verticillate, spread- tane tropical rainforests of the northern two thirds
ing at 4570 from the main branch, retaining leaves of the main island, Grande Terre. It has been found
only towards ends, terete, with obtriangular to oval at altitudes between 500 m and 1350 m a.s.l. and
leaf scars. Terminal buds globose, ca. 3 mm wide and appears on serpentine (ultramafic soils) as well as
23 mm long, with triangular, apiculate scales free on acidic mica schist (Mt. Canala, Mt. Pani) or
at apex. Leaves alternate (helical), on juvenile plants gneiss (Mt. Ignambi). It is associated with other
longer than on mature trees, up to 17 cm long; on conifers and angiosperms commonly found in the
mature plants (4)69(12) cm long, spreading for- south of the island, so its (seeming?) absence there is
ward at 3060 to shoot axis, crowded towards ends somewhat mysterious. Its superficial resemblance to
of branchlets except on vigorous leading shoots, nar- Podocarpus spp., with which it may occur together,
rowly lanceolate, 3.57(9) mm wide near the mid- makes it difficult to detect unless the tree is fertile,
dle, straight or slightly falcate, gradually widening but other things being equal this applies to its entire
from a 58 mm long petiolate base, gradually taper- range.
Conservation serpentine deposits (garnierite) within its altitu-
dinal range.
This interesting species, an endemic to Grande Terre, IUCN: NT
appears to be relatively common and is not specifi-
cally exploited for its timber. It occurs often in inac- Uses
cessible places and at least in several forest reserves,
e.g. Table Unio, Mt. Pani, and Roches de Ouaime No uses have been recorded of this species and it is
(proposed). The quest for anti-cancer drugs found not known to be in cultivation except as a few young
in the alkaloid compounds of the family Taxaceae, plants in perhaps three botanic gardens (BGCI
which led to the exploitation of several species, does database, accessed May 2008). Its wood properties
not seem to have affected this isolated species seri- presumably resemble those of other members of 229
ously. On the other hand, deforestation probably has Taxaceae, which are not commercial timber trees but
had an impact on it, particularly in nickel mining can be put to specialized uses like furniture, wood
concessions in the north of the island, and it is not turning and arts and crafts. This interesting species
safe from further developments in this increasingly should be grown more in botanic gardens and other
spreading activity, particularly aimed at montane tree collections where climatic conditions are suitable.
Callitris Vent., Decas Gen. Nov.: 10. 1808. Frenela Mirb., Mm. Mus. Hist. Nat. 13:
30. 1825. [Frenela triquetra Spach] Type: Callitris rhomboidea R. Br. ex Rich. & A.
Rich. (Cupressaceae).
Octoclinis F. Muell., Trans. & Proc. Philos. Inst. the upper whorl of scales. Seeds moderately numer-
Victoria 2: 21. 1858. Type: Octoclinis macleayana F. ous, with 23 wings. Seedlings with 2 cotyledons.
Muell. [Callitris macleayana (F. Muell.) F. Muell.].
Nothocallitris A. V. Bobrov & Melikyan, Komarovia 15 species.
4: 84. 2006. Type: Nothocallitris sulcata (Parl.) A. V.
230 Bobrov & Melikyan [Callitris sulcata (Parl.) Schltr.]. Distribution
Description Synopsis
Shrubs or trees, evergreen, monoecious. Resin cavi- In the Monograph of Cupressaceae and Sciadopitys
ties in leaves. Bark on trunks of large trees thick, (Farjon, 2005a) two sections were accepted: section
deeply fissured, hard (soft and stringy in one species). Callitris with 14 species and section Octoclinis with a
Branches spreading or ascending, forming a bushy, single species, Callitris macleayana. A phylogenetic
conical, pyramidal or broadly domed crown. analysis based on multiple-gene DNA sequence data
Fastigiate forms common in several species in and comprehensive taxon sampling is still wanting.
nature. Ultimate branchlets covered with leaves, Despite this, the separation of the morphologically
lateral branchlets shed continuously. Leaves scale- and ecologically distinctive species C. macleayana at
like, in alternate whorls of 3 (in one species also with section rank seems warranted and is here accepted.
acicular, spreading leaves in whorls of 4), decurrent,
appressed, linear, abaxially convex or keeled, con- Callitris sect. Callitris
nate but parting with the thickening of branches, Species: Callitris bailey C. T. White, C. canescens
persisting several years, extremely variable in size (Parl.) S. T. Blake, C. columellaris F. Muell., C.
dependent on growth of branches but on ultimate drummondii ((Parl.) F. Muell., C. endlicheri (Parl.)
branchlets mostly less than 5 mm long; margins usu- F. M. Bailey, C. monticola J. Garden, C. muelleri
ally minutely denticulate, apical part slightly differ- (Parl.) Benth. & Hook. f. ex F. Muell., C. neocale-
entiated (e.g. thicker) from main length of the leaf. donica Dummer, C. oblonga Rich. & A. Rich., C.
Pollen cones solitary or with 23 together at the tips preissii Miq., C. rhomboidea R. Br. ex Rich. & A.
of branchlets, small, short cylindrical; microsporo- Rich., C. roei (Endl.) F. Muell., C. sulcata (Parl.)
phylls 820 in whorls of 3 or decussate, with 26 Schltr., C. verrucosa (A. Cunn. ex Endl.) F. Muell.
abaxial pollen sacs. Seed cones terminal on short
leafy shoots, solitary or more often clustered and Callitris sect. Octoclinis (F. Muell.) Benth. & Hook. f.
serotinous, (sub)globose when closed, early decidu- Species: Callitris macleayana (F. Muell.) F. Muell.
ous or more often persistent on branches and stems.
Bract-scale complexes in two alternate whorls of 3 (in Key to the sections of the genus Callitris
one species also in two whorls of 4), with the upper
whorl usually the largest, valvate, thick, smooth, 1a. Seed cones with two whorls of ternate scales,
rugose or with verrucae on the abaxial surface and only on branches with adult scale leaves. Bark
with light seed marks on the adaxial surface; bract smooth or hard and scaly Section Callitris
tips subapical, small. Columella distinct, variably
shaped, often trimerous, the lobes alternating with
1b. Seed cones with two whorls of ternate as well as 9b. Mature seed cones angular-globose or longer
quadrate scales, the latter on branches with than wide, with large subapical bract tip
juvenile needle leaves, both occurring in mature deforming the cone scale 11
trees. Bark soft and stringy Section Octoclinis, 10a. Seed cones (10)1215 mm wide, with tapering
a single species Callitris macleayana base from thickened shoot terminus; leaves
obtusely keeled C. drummondii
Key to the species of section Callitris 10b. Seed cones 1525 mm wide, with abrupt base
from shoot terminus; leaves acutely keeled
1a. Seed cones at the terminal end of thin foliage C. muelleri
branchlets, solitary or a few together, caducous 11a. Seed cones mostly longer than wide, with
2 markedly different lengths of scales in each 231
1b. Seed cones on short, thickened foliage branch- whorl C. oblonga
lets, usually clustered, persistent 6 11b. Seed cones angular-globose; scales more equal
2a. Dorsal surface of adult leaf rounded, not in length 12
keeled 3 12a. Seed cones whitish grey to metallic grey, weath-
2b. Dorsal surface of adult leaf distinctly (obtusely ering to dull grey. Restricted to SW Western
or acutely) keeled 4 Australia C. roei
3a. Shrub or small tree. Seed cone scales narrowly 12b. Seed cones darker coloured. Eastern Australian
oblong and nearly equal in size, ca. 3 mm wide. species 13
Endemic in New Caledonia C. sulcata 13a. Dwarf tree 13(4) m tall; bark remaining
3b. Tree. Seed cone scales unequal in size, wider smooth; leaves on ultimate branchlets 23 mm
than 4 mm. Widespread in mainland Australia long. Restricted to a small area in NE New
C. columellaris South Wales and SE Queensland
4a. Shrub or small tree. Juvenile and transitional C. monticola
leaves persisting on young trees. Endemic to 13b. Shrub or tree to 1015 m; bark soon scaly, fis-
New Caledonia C. neocaledonica sured; leaves on ultimate branchlets 36 mm
4b. Trees. Juvenile and transitional leaves restricted long. Widespread from Queensland to
to seedlings. Native in mainland Australia 5 Tasmania C. rhomboidea
5a. Bark of lower trunk thick, deeply fissured;
leaves obtusely keeled C. endlicheri
5b. Bark on lower trunk thin, shallowly fissured; Callitris baileyi C. T. White, Proc. Linn. Soc.
leaves acutely keeled C. baileyi New South Wales 48: 449. 1923. Type: Australia:
6a. Dorsal surface of adult leaf rounded, not Queensland, Blackbutt Range, Benarkin, W. D.
keeled 7 Francis [BRI 192911] s.n. (lectotype BRI).
6b. Dorsal surface of adult leaf distinctly (obtusely
or acutely) keeled 9 Etymology
7a. Seed cones verrucose when mature, more or
less covered with small or large verrucae 8 This species was named after Frederick M. Bailey
7b. Seed cones smooth or rugose when mature, (18271915) who collected the first specimens.
verrucae absent C. canescens
8a. Verrucae variable in density, often quite large in Vernacular names
proportion to the cone, sometimes nearly
absent C. preissii Baileys Cypress-pine
8b. Verrucae dense, equally distributed and never
nearly absent, small in proportion to the cone Description
C. verrucosa
9a. Mature seed cones globose, smooth, with very Trees to 1518 m tall, often less than 10 m; monopo-
small subapical bract tip not deforming the dial, usually branching low and often fastigiate, up to
cone scale 10 50 cm d.b.h. Bark on trunk thin, shallowly furrowed
or fissured, hard, slowly exfoliating in small flakes Ecology
and strips, greyish, inner layers light brown.
Branches ascending at ca. 45 or steeper, slender, In Eucalyptus forest or woodland, with Eucalyptus
forming a conical or pyramidal, often symmetrical crebra or other species, often in small groups scat-
crown. Foliage branchlets sparse, ascending or erect, tered through forest; in hills and low mountains on
twisted and rigid, triangular in cross section, 0.81.2 basalt and volcanic ash or in sandy soil with boul-
mm diam., covered with closely appressed leaves. ders. The climate is mild, with most rainfall in the
Leaves in alternate whorls of 3, decurrent, closely summer.
appressed, apices appressed or free on some lead-
ing shoots (whip shoots), connate but parting with Conservation
232 thickening of branches, persisting several years, lin-
ear, 25(7) mm long on ultimate branchlets, 0.71.2 Decline has mainly come from changes in habitat
mm wide, abaxially acutely keeled; margins den- due to pressures from grazing and increased hazard
ticulate near obtuse or acute apex; epistomatic, sto- of fires. Several populations are now within pro-
mata in two narrow, marginal bands; abaxial surface tected areas (National Parks), but decline is likely to
smooth, green. Pollen cones solitary or 23 together continue outside these localities.
and terminal on ultimate branchlets, oblong, 23 IUCN: NT
11.2 mm, yellowish green turning light brown;
microsporophylls 814, in alternate whorls of 3 or Uses
decussate near apex, peltate, with hyaline, erose-
denticulate margins, bearing 23 abaxial pollen The wood of this species was traditionally used for
sacs near the lower margin. Seed cones terminal fence posts. It is extremely rare in cultivation, lim-
on short, thin, leafy shoots, solitary, maturing in ited to a few botanic gardens.
11.5 years, caducous, subglobose to broadly ovoid
when closed, 1013 912 mm, smooth but becom-
ing rugose when open, brown or purplish brown Callitris canescens (Parl.) S. T. Blake, Proc. Roy.
weathering grey. Bract-scale complexes in 2 alter- Soc. Queensland 70: 39. 1959. Frenela canescens
nate whorls of 3, deeply incised towards cone base Parl. in Candolle, Prodr. 16 (2): 448. 1868. Type:
but not opening very wide; upper ones largest, ca. Australia: Western Australia, [Int. S.W. Australia],
10 5 mm, tapering to an obtuse apex; smaller ones J. S. Roe s.n. (lectotype G?, n.v., isolectotype K).
narrowly triangular, acute; bracts largely included, Fig. 61
showing a prominent apex below the apex of each
scale; adaxial surface reddish brown or brown with Etymology
light seed marks. Columella robust in comparison to
small cone (as high as shortest scales in open cones), The species epithet means greyish or turning grey
34 22.5 mm, more or less triangular in cross- and may refer to the seed cones.
section, usually slightly narrowed at base, obtuse.
Seeds 24(5) on each scale, ovoid or triangular, Vernacular names
flattened, up to 67 mm including wings (seed body
23 mm), brown with large, whitish concave hilum, Morrisons Cypress-pine
wings 2, lateral, very unequal in size and shape, up to
4 mm long. Description
Ecology Etymology
In dwarf scrubland (kwongan) with Myrtaceae The species epithet refers to a columella, i.e. the cen-
(Melaleuca), Proteaceae (e.g. Banksia, Dryandra, tral, columnar structure in the seed cone.
Grevillea, Hakea), or low Eucalyptus woodland
(mallee) with other Myrtaceae (e.g. Melaleuca), Vernacular names
Proteaceae (e.g. Banksia, Grevillea, Hakea), Acacia
spp., and Allocasuarina spp.; often in strips of natural White Cypress-pine, White Pine, Murray River
vegetation between roads and ploughed fields or in Pine, Murray Pine, Western Sand Cypress, Western
disturbed vegetation; on plains or low ridges (break- Cypress, Cypress Pine, Northern Cypress-pine
Description Taxonomic notes
Trees to 20 m tall; trunk monopodial, usually Treatment of Callitris glaucophylla and C. intra-
branching low or occasionally forked, up to 50 cm tropica as taxonomic synonyms under a (slightly)
d.b.h. Bark soon scaly, on trunk deeply fissured, broader circumscription of C. columellaris has been
hard, exfoliating in small flakes and strips, grey- explained and justified in detail in the Monograph of
brown. Branches spreading or near the top ascend- Cupressaceae and Sciadopitys (Farjon, 2005a).
ing, long, forming a conical, pyramidal or broadly
domed, more or less open crown. Foliage branch- Distribution
lets numerous, spreading or ascending, ultimately
234 very slender, terete, 0.71.0 mm diam., covered Australia: In all states and territories except
with closely appressed leaves, persistent. Leaves in Tasmania, but absent from low lying deserts and
alternate whorls of 3, decurrent, closely appressed, moist coastal regions.
apices appressed or free on some leading shoots TDWG codes: 50 NSW-NS NTA QLD-QU SOA VIC
(whip shoots), connate but parting with thicken- WAU-WA
ing of branches, persisting several years, linear, 26
mm long on ultimate branchlets, 0.40.8 mm wide, Ecology
abaxially convex; margins denticulate near slightly
broadened acute-acuminate apex; epistomatic, sto- In a wide range of semi-arid to more mesic habitats,
mata in two marginal lines, abaxial stomata near from banks or dunes near the sea to rocky canyon
base of leaves only or absent; abaxial surface weakly bottoms and sandstone plateaux in the interior. In
verrucose, green or glaucous, variable with many open Eucalyptus forest or low woodland (mallee)
intermediate shades of colour. Pollen cones numer- with Eucalyptus spp., Melaleuca spp., Allocasuarina
ous, solitary or paired and terminal on ultimate spp., Acacia spp., Banksia spp., Grevillea spp.,
branchlets, ovoid-oblong, 34 1.5 mm, yellowish Triodia spp., and Callitris endlicheri; in disturbed
green turning orange-brown; microsporophylls (grazed) open woodland; in the dry interior in
1018, in alternate whorls of 3, peltate, with erose Acacia low woodland or scrub (mulga) with A.
margins, bearing 3(4) abaxial pollen sacs near the aneura and other Acacia spp., Eremophila spp., in
lower margin. Seed cones terminal on short, thin, hummock grassland/ scrub with spinifex (Triodia
leafy shoots, solitary but often numerous on higher sp.); and in rocky (sandstone) canyons and on ridges
order branches, maturing in 11.5 years, caducous, with Eucalyptus spp., Acacia spp., and Macrozamia
globose when closed, (7) 1018 (10)1220 mm, macdonnellii. On (coastal) sand dunes near sea level,
smooth or rugose to slightly pustulate, glaucous- on plains, hills, tablelands, ridges and low moun-
brown or purplish brown weathering grey. Bract- tains to 1300 m a.s.l. in white, yellow, brown or red
scale complexes in 2 alternate whorls of 3, deeply sand or loam or on gravelly laterite (hardpan) over
incised towards cone base when opened; upper ones sandstone, ironstone, conglomerate, gypsum, shale,
largest, 812 68 mm, tapering to an obtuse apex; basalt or granite. The climate varies from mesic and
smaller ones narrower 68 46 mm, acute; bracts mild on the coast of New South Wales to arid in the
almost entirely included, showing a minute prickle interior of Australia; for most of its range this species
below the apex of each scale; adaxial surfaces red- occurs within the zone with summer rainfall.
dish brown or dark brown with light seed marks
towards base. Columella variably shaped, up to 6 Conservation
mm long, obtuse or acute-triangular. Seeds 410
on each scale (the higer number on larger scales), IUCN: LC
triangular, flattened, up to 8 mm including wings
(seed body up to 45 23 mm), dark brown Uses
with whitish concave hilum; wings 2 on opposite
sides, 45 mm wide, more or less equal in shape Timber of this species was (is?) used for panelling in
and size. cabinet making, and also for fence posts when these
were still predominantly of wood. As with many lines; abaxial surface weakly verrucose or smooth,
species in Cupressaceae, the wood is resistant to yellowish green or green. Pollen cones solitary or in
rot and, in this case, also to termite attack. Baker & pairs and terminal on ultimate branchlets, ovoid-
Smith (1910) reported a decline in its availability in oblong, 36 1.52 mm, yellowish green turning
many regions without efforts to restock it. Its exploi- light brown; microsporophylls 1220, proximal ones
tation is now much less intensive and regeneration is in alternate whorls of 3, distal ones decussate, sub-
taking place in many localities. Its horticultural use peltate, with hyaline-denticulate margins; apex acu-
is very limited. In Australia, there seems to be little minate; bearing 3 abaxial pollen sacs near the lower
interest in this and other native conifers, but its vari- margin. Seed cones terminal on short, 56 mm thick,
ability in foliage colour from dark green to extremely leafy shoots, solitary but often grouped along limited
glaucous should afford scope for horticultural selec- lengths of stems, maturing in 11.5 years, persistent, 235
tion and planting in regions with a Mediterranean or (depressed-)globose when closed, tapering at base to
even desert climate. thickened terminus of shoot, (10)1215 (10)1217
mm, smooth or rugose when dry with opened scales,
purplish grey to metallic grey. Bract-scale complexes
Callitris drummondii (Parl.) F. Muell., Syst. Census in 2 alternate whorls of 3, not deeply incised towards
Austral. Pl. 1: 109. 1882. Frenela drummondii Parl. cone base and hence not opening widely; upper ones
in Seemann, J. Bot. 1: 35. 1863. Type: Australia: largest, 1012 810 mm, tapering to an angular-
Western Australia, Southern Coastal Region, [In obtuse apex; smaller ones narrower 810 68 mm,
Nova Hollandia austro-occidentali ad Cynorem acute, all very thick; bract tip minute, only visible in
flumen legit Cl. J Drummond], J. Drummond smooth, closed cones below the apex of each scale;
[ex herb. F. von Mueller] 433 (holotype FI). adaxial surfaces dark brown with light seed marks
at margins near base. Columella very thick, 35 mm
Etymology long and wide at base, (tri-)angular or terete, obtuse.
Seeds usually only 2 on each scale, angular, flat-
This species was named after J. Drummond, who tened, up to 7 mm including wings (seed body 23
collected the type specimens. 2 mm), brown with whitish concave hilum; wings 2
on opposite sides, 12 and 24 mm wide, unequal in
Vernacular names shape and size.
plate 8. Callitris monticola. 1. Habit of trees. 2. Foliage branch. 3. Branchlet with juvenile and adult
leaves. 4. Pollen cone. 5. Shoot with pollen cones and seed cones. 6. Cluster of seed cones. 7. Seeds.
Description Distribution
Dwarf trees 13(4) m, sometimes a shrub, monoe- Australia: SE Queensland, NE New South Wales,
cious; trunk monopodial or multistemmed, branch- from ca. 28S to ca. 30S straddling the Great
ing low, up to 10 cm d.b.h. Bark smooth to slightly Dividing Range.
scaly, thin, slowly exfoliating in small flakes, grey. TDWG codes: 50 NSW-NS QLD-QU
Branches spreading or ascending at ca. 45, long
and pliant, forming an open, irregular or pyramidal Ecology
crown. Foliage branchlets sparse, spreading irregu-
larly or more or less erect, arranged in more or less This species occurs in low Eucalyptus woodland or
240 planated sprays towards ends of branches; ultimate forest and is virtually restricted to rock outcrops and
branchlets very slender, triangular in cross section, cliff edges thinly covered with scrub. Associated with
0.60.8 mm diam., covered with closely appressed Eucalyptus spp., Allocasuarina rigida, Xanthorrhoea
leaves, persistent. Leaves in alternate whorls of 3, sp., Banksia spp., Leptospermum spp., Mirbelia con-
decurrent, closely appressed, apices appressed or fertifolia, and Restio stenocoleus; in very shallow soil
some free especially on some leading shoots (whip amongst boulders or in crevices in granitic rock
shoots), connate but parting with thickening of (Adamellite), trachyte, or sandstone. The altitudinal
branches, persisting several years, linear, 23 mm range is from 560 m to 1360 m a.s.l. The climate is
long on ultimate branchlets, 0.50.6 mm wide, mild and mesic, with a peak of rainfall during the
abaxially prominently keeled; margins denticulate summer.
near slightly broadened acute apex; epistomatic, sto-
mata mainly in two narrow marginal bands; abaxial Conservation
surface densely verrucose, yellowish green to glau-
cous. Pollen cones solitary and terminal on ultimate This species has a restricted distribution with very
branchlets, ovoid-globose, 2.5 2 mm, yellowish localized populations in a vegetation type that is
green turning orange-brown; microsporophylls under increased threat of fires. On one locality
912, in alternate whorls of 3, peltate, apiculate, with (summit area of Bald Rock) it could not be found
erose-denticulate margins, bearing 23 abaxial pol- again despite a thorough search in 1997 by me and
len sacs near the lower margin. Seed cones termi- earlier searches by J. Williams from Armidale; only
nal on short, thick, leafy shoots, mostly aggregated the common C. rhomboidea remains there.
in dense, compact clusters on stems and branches, IUCN: EN [B2ab (iv)]
maturing in 11.5 years, persistent, subglobose when
closed, 1220 1218 mm, verrucose or nearly Uses
smooth, slightly rugose when open, dull brown,
weathering dull grey. Bract-scale complexes in 2 No uses are known of this species.
alternate whorls of 3, not deeply incised towards base
and opening only slightly; upper ones largest, 1016
610 mm, with parallel or slightly widening sides Callitris muelleri (Parl.) Benth. & Hook. f. ex
and obtuse apex; smaller ones wider at base, triangu- F. Muell., Syst. Census Austral. Pl. 1: 109. 1882.
lar, with acute apex; bract tip forming a small, acute Frenela muelleri Parl., in Candolle, Prodr. 16 (2):
umbo below the apex of each scale; abaxial surface 450. 1868. Type: Australia: New South Wales,
convex, brown turning grey, with conspicuous white Sydney, Port Jackson, [South Head], A. Thozet
(dotted) margins; adaxial surface dark brown with [ex herb. F. von Mueller] MEL 503745 (lectotype
inconspicuous seed marks. Columella large, mostly MEL). Fig. 65, 66
trilobed, up to 8 mm wide, obtuse. Seeds 35 on
basal part of each scale, more or less triangular, flat- Etymology
tened, up to 8 mm including wings (seed body 45
23 mm), dark brown, with concave hilum; wings 2 This species was named after F. J. H. von Mueller
on each side, up to 4 mm wide, unequal or nearly (18251896), the famous German botanist working
equal in shape and size. on the Australian flora.
Vernacular names up to 5 3 mm, dark brown with lighter coloured
concave hilum; wings 2, 25 mm wide, more or less
Illawarra Cypress-pine, Illawarra Pine equal in shape and size.
Description Distribution
Dwarfed trees or shrubs to 34.5(6) m; monopo- Australia: E-central New South Wales, from areas
dial or multistemmed, branching low and often near the coast to the table lands of the Blue Mts. E
fastigiate, up to 15 cm d.b.h. Bark smooth, not fis- of Sydney.
sured, thin, slowly exfoliating in small flakes and TDWG codes: 50 NSW-NS
strips, grey. Branches ascending at ca. 45 or erect, 241
short or long in fastigiate crowns, slender, forming Ecology
a conical to pyramidal, in fastigiate forms columnar
crown. Foliage branchlets numerous, in dense tufts In scrub vegetation associated with Myrtaceae (e.g.
towards end of main branches, ultimately slender, Calytrix, Eucalyptus apiculata, E. stricta, E. sieberi,
triangular in cross section, 0.71 mm diam., covered Leptospermum), Proteaceae (e.g. Banksia, Grevillea),
with closely appressed leaves, persistent. Leaves in Allocasuarina distyla, Lomandra glauca, and Caustis
alternate whorls of 3, decurrent, closely appressed, pentandra. On skeletal sandy or gravelly soil and
apices appressed or free on some leading shoots among boulders or in rock crevices of sandstone on
(whip shoots), connate but parting with thicken- cliffs or talus slopes, sometimes congregating on dirt
ing of branches, persisting several years, linear, 28 road verges. The altitude ranges from 500 m to 1050
mm long on ultimate branchlets, 0.50.7 mm wide, m a.s.l. The climate is temperate and mesic and frost
abaxially keeled; margins denticulate near slightly can occur in winter at the higher altitudes.
broadened obtuse or sometimes acute apex; episto-
matic, stomata in two narrow, marginal bands; abax- Conservation
ial surface minutely verrucose, green. Pollen cones
numerous, solitary or with 23 together, terminal on Some of the earlier populations reported from the
ultimate branchlets, ovoid-globose to oblong, 23 Sydney (Port Jackson) area have undoubtedly been
11.5 mm, yellowish green turning orange-brown; reduced in area of occupancy due to urbanisation.
microsporophylls 912, in alternate whorls of 3, pel- Fires, if too frequent, will threaten regeneration suc-
tate, with denticulate margins, bearing 23 abaxial cess. Away from the urbanised areas the species,
pollen sacs near the lower margin. Seed cones ter- though not widespread, seems to hold its own in sites
minal on short, thick, leafy shoots, solitary but often that support no commercial land use of any kind.
grouped together in small clusters on branches and IUCN: NT
stems, maturing in 11.5 years, persistent, subglobose
or globose when closed, 1525 1525 mm, smooth Uses
and lustrous when closed, coarsely rugose when
open, greenish or glaucous when growing, maturing There are no economic uses reported of this species.
to brown or purplish brown weathering grey. Bract-
scale complexes in 2 alternate whorls of 3, very thick,
not deeply incised towards cone base when opened; Callitris neocaledonica Dummer, J. Bot. 52: 239.
upper ones much larger than the lower, to 20 13 1914. Nothocallitris neocaledonica (Dummer) A. V.
mm, usually widest above the middle and tapering to Bobrov & Melikyan, Komarovia 4: 86. 2006. Type:
an obtuse apex; smaller ones narrower and tapering New Caledonia: Grande Terre, Province Sud, Ngoye
to an acute apex; a minute prickle (bract tip) below River, [auf den Bergen am Ngoye], R. Schlechter
the apex of each scale; adaxial surface red-brown 15179 (holotype K).
or dark brown with numerous whitish seed marks.
Columella triangular, sometimes flattened, ca. 3 mm Etymology
long and wide, acute, dark brown to black. Seeds
510 on each scale, ovoid or triangular, flattened, The species epithet refers to its country of origin.
Vernacular names scale, ovoid or triangular, flattened, 57 22.5 mm,
light brown, with small hilum; wings 2 on each side,
Ni up to 7 1.5 mm, unequal in shape and size.
Description Distribution
Shrubs or small trees 37(10) m tall; trunk monopo- New Caledonia, Grande Terre, Province Sud.
dial or very short, branching low, up to 50 cm diam. TDWG codes: 60 NWC
at base. Bark soon scaly, on lower trunk becoming
shallowly fissured, rough, exfoliating in long strips, Ecology
242 brown weathering grey. Branches spreading and
assurgent, thick and long, often contorted, form- In scrub vegetation (maquis minier) or low sclero-
ing a sympodial, rounded and open crown. Foliage phyll forest on slopes or along creeks, in shallow soil
branchlets in dense tufts at ends of main branches, over ultramafic (serpentine) rocks. The altitudinal
assurgent or erect, ultimate branchlets short, rigid, range is between 560 m and 1500 m a.s.l. The climate
thick, apprearing conspicuously articulate with scale is low montane tropical with high rainfall through-
leaves, triangular in cross section, 1.52 mm diam., out the year.
covered with closely appressed leaves, green, persis-
tent. Leaves of two types on plants up to 1 m tall. Conservation
Mature (scale) leaves (and transitional leaves) in
alternate whorls of 3, decurrent, closely appressed, This species has a limited distribution in localized
apices appressed or some free especially on some populations in the southern massifs. It is protected
leading shoots (whip shoots), connate but part- in the Montagnes des Sources and Mt. Humboldt
ing with thickening of branches, linear, 2.55 mm reserves, but fire is a hazard and adequate manage-
long on ultimate branchlets, ca. 1 mm wide, acutely ment of this increasing problem is urgently needed.
keeled abaxially; margins erose-denticulate near IUCN: NT
obtuse or acute apex; epistomatic, stomata in two
narrow marginal bands; abaxial surface smooth Uses
on keel, minutely verrucose on sides, green. Pollen
cones solitary and terminal on ultimate branchlets, There are no commercial uses recorded of this
ovoid or subglobose, 23 22.5 mm, yellowish species.
turning brown; microsporophylls 912, in alternate
whorls of 3, peltate, with erose-denticulate margins,
obtuse-acuminate, bearing 34 abaxial pollen sacs Callitris oblonga Rich. & A. Rich., in A. Richard
near the lower margin. Seed cones few, terminal on (ed.) Comm. Bot. Conif. Cycad.: 49. 1826. Type:
short, thin, scale-leaved shoots, solitary, maturing in Australia: Tasmania, [Nouvelle Hollande], J. L. C.
1 year, caducous (or persistent a short time after seed Dumont dUrville s.n. (holotype P).
dispersal), ovoid when closed, 810 67 mm, with
convex scales when open, smooth, greenish matur- Callitris oblonga Rich. & A. Rich. subsp. corangensis
ing to brown. Bract-scale complexes in two alternate K. D. Hill, Fl. Australia 48: 716. 1998.
whorls of 3, not opening very wide, nearly equal in Callitris oblonga Rich. & A. Rich. subsp. parva K. D.
size and shape, 78 3 mm, narrowly oblong, slightly Hill, Fl. Australia 48: 717. 1998.
tapering, rectangular in cross-section, with mostly
included bracts forming a prominent and acute Etymology
umbo below the apex of each scale; adaxial surface
smooth, red-brown with inconspicuous seed marks. The species epithet refers to the oblong (longer than
Columella very long, 56 1.52 mm, angular, with wide) seed cones.
triangular apex. Seeds 1(2) at base of each upper
Vernacular names dark brown to black with large, yellowish concave
hilum; wings 2, lateral, 34 mm wide, more or less
Tasmanian Cypress-pine, Dwarf Cypress-pine, equal in shape and size.
Native Cypress, River Pine, Pigmy Cypress-pine
Taxonomic notes
Description
For a discussion of the two subspecies described in
Shrubs or small trees to 78 m tall, often less than 4 Flora of Australia 48 (1998), here treated as taxo-
m; monopodial or multistemed from a short basal nomic synonyms of Callitris oblonga, the reader
trunk, usually branching low and often fastigiate, is referred to the Monograph of Cupressaceae and
up to 20 cm d.b.h. Bark soon scaly, on the lower Sciadopitys (Farjon, 2005a). 243
trunk of largest trees fissured, hard, slowly exfoliat-
ing in small flakes and strips, grey-brown. Branches Distribution
ascending or erect, long in fastigiate crowns, slen-
der, forming a conical or pyramidal, open or dense Australia: New South Wales, in two disjunct areas:
crown, or a more or less rounded shrub in less favor- in the New England Tablelands and adjacent moun-
able sites. Foliage branchlets ascending to erect, tains, and along the Corang River, a tributary of the
ultimate branchlets triangular in cross section, 11.2 Shoalhaven River; and in NE Tasmania along several
mm diam., covered with closely appressed leaves, rivers.
persistent. Leaves in alternate whorls of 3, decurrent, TDWG codes: 50 NSW-NS TAS
closely appressed, apices appressed or free on some
leading shoots (whip shoots), connate but part- Ecology
ing with thickening of branches, persisting several
years, linear, 312 mm long on ultimate branchlets, Mostly riparian, in Eucalyptus woods or shrubland;
0.71 mm wide, abaxially keeled; margins denticu- in Tasmania often in disturbed vegetation with
late near slightly broadened acute-acuminate apex; invasives like Ulex europaeus, Rubus discolor, and
epistomatic, stomata in two narrow, marginal bands; Crataegus sp. Associated taxa are Eucalyptus spp.,
abaxial surface weakly verrucose, green or glaucous Acacia spp., Leptospermum spp., Allocasuarina spp.,
green. Pollen cones subterminal in clusters of (2)3 Bursaria spinosa, Callistemon paludosis, Jacksonia
5(6) on ultimate branchlets, ovoid-globose, 11.5 1 scoparia, Lomandra longifolia, Epacris microphylla,
mm, yellowish green turning orange-brown; micro- Hakea spp., Lomatia myrtifolia, Pteridium sp., and
sporophylls 812, in alternate whorls of 3, peltate, grasses (Poaceae). The altitudinal range is from 10
rounded, with entire margins, bearing 23 abaxial m to 1300 m a.s.l., with the Tasmanian populations
pollen sacs near the lower margin. Seed cones termi- mostly below 100 m a.s.l. Soils are (gravelly) sand or
nal on very short, thick, leafy shoots, rarely solitary loam, often rocky with granitic boulders or bedrock,
and often grouped together in compact clusters on acidic, and sometimes silt or clay, subject to regu-
branches and stems, maturing in 11.5 years, persis- lar flooding. In a few cases in Tasmania, C. oblonga
tent, ovoid-oblong or ovoid-utriculate when closed, appears to occur away from streams on floodplains,
(12)1520(25) (10)1216(20) mm, smooth, invariably on disturbed sites grazed by cattle, and in
rugose when dry, blackish brown or black, weather- New South Wales on moderate slopes with seepage
ing grey. Bract-scale complexes in 2 alternate whorls water. The climate is temperate and mesic, without
of 3, deeply incised towards cone base when opened, prolonged periods of drought, but with higher rain-
thick, parting only slightly; upper ones largest, up to fall levels in the uplands of New South Wales than in
20 10 mm, tapering to a (recurved) obtuse apex; NE Tasmania.
smaller ones 1/2 to 3/5 the size of the larger ones,
acute or obtuse; adaxial surfaces dark purplish Conservation
brown with light seed marks. Columella small, tri-
angular to trilobed, 23 mm long, obtuse. Seeds 515 Briggs & Leigh (1988) list Callitris oblonga (N.S.W.)
on each scale, ovoid or triangular, flattened, up to and Callitris sp. 1 [= C. oblonga in Tasmania] as
10 mm including wings (seed body 34 23 mm), Vulnerable (3VCa in CSIRO conservation coding).
More recent research (Nadolny & Benson, 1993) has Etymology
confirmed a wider occurrence in the New England
Range of N.S.W. than was previously known. Some This species was named after Ludovicus (Ludwig)
of the populations are in conservation areas, but Preiss, who collected this species on Rottnest Island
most are outside these on rangeland or bushland. in 1839.
Threats are conversion to agriculture by clearance of
natural vegetation (in particular Corang River and Vernacular names
Tasmania), grazing of livestock, associated invasive
species (especially Ulex europaeus) and fires if too Rottnest Island Pine, Mallee Pine, Murray Pine,
frequent (with intervals of 3 years or less). Some Mountain Pine, Mountain Cypress-pine, Lachlan
244 populations are (now) very small and isolated. In Pine, Light Pine, White Pine, Common Cypress-pine
Tasmania a recovery programme is under way using
several approaches. Description
IUCN: VU [B1ab (iii)]
Shrubs or trees to 25 m tall; monopodial or multi-
Uses stemmed, usually branching low and frequently
forked, up to 1 m d.b.h. Bark soon scaly, on large
There is a limited ornamental use of this species in trunks thick, deeply fissured, stringy, exfoliating in
Tasmania. In cultivation, as with many species in large strips, light brown; outer bark grey. Branches
Cupressaceae, Tasmanian Cypress-pine grows con- spreading or ascending, long, forming a pyrami-
sistently in a fastigiate habit. In the wild, this habit dal or broadly domed, more or less open crown,
is commonly found in Tasmania but, as far as I have or a pyramidal or rounded shrub. Foliage branch-
seen, absent in New South Wales. It is possible that lets spreading or ascending, ultimately very slen-
there is a genetic basis for this, but environmental der, terete, 0.71.0 mm diam., covered with closely
factors also seem to play a role, although it remains appressed leaves, persistent. Leaves in alternate
unclear what these are specifically. From what I have whorls of 3, decurrent, closely appressed (except on
seen in this species in both areas, it could be that some whip shoots), connate but parting with thick-
more adverse conditions for fast growth prevent ening of branches, persisting several years, linear,
fastigiate branching and promote spreading branch- 1.55 mm long on ultimate branchlets, 0.51 mm
ing instead. The fastigiate form should be cultivated wide, abaxially convex; margins denticulate near
more widely, as it forms a well shaped erect shrub or obtuse or acute apex; epistomatic, a few abaxial sto-
small tree. mata near base of leaves only or absent, adaxial sto-
mata in two marginal bands and a few below apex;
abaxial surface verrucose, yellowish green or some-
Callitris preissii Miq., in Lehmann, Pl. Preiss. 1: 643. times glaucous green. Pollen cones solitary or 23
1845. Type: Australia: Western Australia, Rottnest together, terminal on ultimate branchlets, ovoid-
Island, C. A. Gardner 175 (epitype PERTH), L. Preiss oblong, 35 1.52.5 mm, yellowish green turning
1310 (lectotype L). Fig. 67 orange-brown; microsporophylls 1520, in alternate
whorls of 3 or decussate, peltate, ovate-orbicular,
Callitris tuberculata R. Br. ex R. T. Baker & H. G. with entire or erose margins, bearing 24 abaxial
Smith, Res. Pines Austral.: 99. 1910. pollen sacs near the lower margin. Seed cones ter-
Callitris preissii Miq. subsp. murrayensis J. Garden, minal on short, thick, leafy shoots, solitary or more
Contr. New South Wales Natl. Herb. 2 (5): 373. 1957; often clustered on branches and stems, matur-
Callitris preissii Miq. var. murrayensis (J. Garden) ing in 11.5 years, persistent, (depressed-)globose
Silba, Phytologia Mem. 7: 17. 1984; Callitris gracilis to ovoid-globose when closed, (15)2030(35)
R. T. Baker subsp. murrayensis (J. Garden) K. D. Hill, (18)2235 mm, nearly smooth to coarsely rugose
Fl. Australia 48: 716. 1998. and variably covered with large verrucae, brown or
purplish brown to blackish, weathering grey. Bract- Distribution
scale complexes in 2 alternate whorls of 3, not deeply
incised towards cone base when opened, very thick; Australia: New South Wales, South Australia,
the upper ones largest, 1230 813 mm, tapering to Victoria, Western Australia.
an obtuse apex; the smaller ones narrower, acute or TDWG codes: 50 NSW-NS SOA VIC WAU-WA
obtuse; adaxial surfaces grey-brown or brown with
light seed marks. Columella short, thick, up to 5 mm Ecology
long and as wide at base, obtuse or acute-triangu-
lar. Seeds (7)912(16) on each scale, triangular to In coastal and inland dunes, as well as on sandy
ovoid, flattened, up to 10 mm including wings (seed floodplains, hillsides, low ridges (breakaways)
body up to 35 23 mm), dark brown or red-brown and laterite (hardpan). In coastal dune areas it may 245
with whitish concave hilum; wings 2(3) on opposite form more or less dense stands excluding other
sides, 24 mm wide, more or less equal or unequal in trees and shrubs, or it associates with Acacia sp.
shape and size. and Melaleuca lanceolata; elsewhere it occurs in
low Eucalyptus woodland (mallee), associated with
Taxonomic notes other Myrtaceae (e.g. Melaleuca, Leptospermum),
Proteaceae (e.g. Banksia, Dryandra, Grevillea,
Flora of Australia 48 (1998) took a narrow view of Hakea), Acacia spp., Allocasuarina sp., Dodonaea
C. preissii and included only the populations on sp., and Callitris spp.; in (dwarf) scrubland (kwon-
Rottnest Island and around Perth in it. The Flora gan in SW Australia) with Myrtaceae (Melaleuca
accepted C. tuberculata as a widespread species thicket), Proteaceae, and Xanthorrhoea thorntonii; in
occurring along the coast and in the interior of inland dunes and dune slacks with Eucalyptus spp.,
Western Australia and C. gracilis, including subsp. Acacia spp., Myrtaceae, and Callitris verrucosa; on
murrayensis, occurring in South Australia, Victoria sandstone hillsides and ridges with Eucalyptus spp.,
and New South Wales. Investigation of populations Acacia spp., Allocasuarina cristata, Dodonaea sp.,
on sand dunes at Bremer Bay, Western Australia Callitris columellaris, C. rhomboidea, and C. endli-
(which according to Flora of Australia taxonomy cheri. The altitude ranges from near sea level to 670
belong to C. tuberculata), and populations in similar m a.s.l. Soils vary from calcareous, humus-rich old
habitat in two of the classical localities of C. preis- coastal or riparian sand dunes, (where it develops to
sii (Garden Island and Woodman Point) by me in a monopodial or multi-stemmed tree) to white, yel-
1997 revealed that they are the same. Multistemmed low, brown or red sand or loam flats over limestone,
shrubs are the dominant growth form at Woodman sandstone or laterite (hardpan; on this it develops
Point and common at Bremer Bay and on Garden only to a medium height shrub); also in disturbed
Island. The wartiness of cones as well as their size are road verges through arable land or pasture. The cli-
quite variable in all these populations. The plants in mate is mild to hot and characterized by dry periods
the interior are more distinct, especially those that (which can be extensive in the interior parts of the
grow on lateritic soil types, being (smaller) shrubs, range) and winter rainfall.
never a tree, and having smaller cones. In Victoria,
I observed a similar situation: on sand dunes in Conservation
the Murray River Basin and in the Little and Big
Deserts I found mostly trees, but in the hardpan IUCN: LC
areas between the dunes I found shrubs. Cones are
less densely warty here than in Western Australia, Uses
but considering its variation in that state, I think this
is insufficient evidence to separate C. gracilis from The larger trees with monopodial growth that occur
C. preissii and a broad concept of the species is here locally have been used for timber in local construc-
retained. Garden (1957) further confounded the tax- tion, but its economic value is now limited and no
onomy of this species by postulating two naturally commercial harvest occurs. It is only known in
occurring hybrids. horticulture from arboreta and botanic gardens in
countries with mild winters, but it can endure light pollen sacs near the lower margin. Seed cones ter-
frost without much permanent damage. Its attrac- minal on short, thick, leafy shoots, solitary or more
tive, soft foliage makes it a desirable small tree and it commonly aggregated in dense, compact clusters
should be cultivated more often. on branches, maturing in 11.5 years, persistent,
depressed-globose when closed, 1215 1317 mm,
angular with large, protruding umbos, smooth, often
Callitris rhomboidea R. Br. ex Rich. & A. Rich., lustrous, becoming finely rugose when open, black-
in A. Richard (ed.) Comm. Bot. Conif. Cycad.: 47. ish brown or nearly black, weathering grey. Bract-
1826. Type: Australia: New South Wales, Sydney, scale complexes in 2 alternate whorls of 3, only partly
Port Jackson, R. Brown 3107 (holotype BM). Fig. 68 incised towards cone base when opened; upper ones
246 largest, 1013 812 mm, tapering to an obtuse apex;
Etymology smaller ones narrower 68 57 mm; forming a
prominent and acute umbo below the apex of each
The species epithet refers to the rhomboid shape of scale; adaxial surface dark purplish brown with light
the seed cone scales. seed marks. Columella trilobed to tripartite, 38 mm
wide, obtuse. Seeds 410 on each scale, ovoid or tri-
Vernacular names angular, flattened, up to 8 mm including wings (seed
body ca. 3 2 mm), dark brown to black with large,
Port Jackson Pine, Oyster Bay Pine, Cypress-pine, greyish concave hilum; wings 23 on sides, 23 mm
Illawarra Mountain Pine wide, more or less equal in shape and size.
Description Distribution
Shrubs or trees to 1015 m tall; multistemmed or Australia: New South Wales, Queensland, South
monopodial, usually branching low and often fas- Australia, Tasmania, Victoria.
tigiate, up to 40 cm d.b.h. Bark soon scaly, on the TDWG codes: 50 NSW-NS QLD-QU SOA TAS VIC
lower trunk shallowly fissured, hard, slowly exfoli-
ating in small flakes, dark grey-brown, inner layers Ecology
brown. Branches spreading or ascending at ca. 45,
or fastigiate, long and slender, forming a conical Commonly in open woodland or scrubland on
or (broad) pyramidal, dense or more or less open tablelands or mountains, locally on riparian or
crown. Foliage branchlets spreading or ascend- coastal sands, often amongst boulders; associ-
ing to erect, ultimately very slender, triangular in ated with Eucalyptus spp., other Myrtaceae (e.g.
cross section, 0.61 mm diam., covered with closely Kunzea, Leptospermum), Proteaceae (e.g. Banksia,
appressed leaves, persistent. Leaves in alternate Conospermum, Hakea), Acacia spp., Allocasuarina
whorls of 3, decurrent, closely appressed, apices spp., Daviesia ulicina, Dilwenia sp., Leucopogon spp.,
appressed or free especially on some leading shoots Prostanthera lasianthos, Spyridium sp., Hibbertia sp.,
(whip shoots), connate but parting with thicken- Xanthorrhoea sp., Restio sp., Pteridium sp., Callitris
ing of branches, persisting several years, linear, 36 endlicheri, and C. preissii. Soils are (skeletal) white
mm long on ultimate branchlets, 0.30.7 mm wide, or grey sand over sandstone, limestone, shale, gran-
abaxially keeled; margins denticulate near slightly ite (Adamellite), or it is growing in talus, rocky
broadened acute-acuminate apex; epistomatic, sto- creek bottoms, or crevices of bedrock; also in fields
mata in two narrow, marginal bands; abaxial surface (sheep pasture), with invasives (e.g. Ulex europaeus
weakly verrucose, yellowish green to glaucous green. in Tasmania) as indicators of disturbance. The altitu-
Pollen cones solitary or paired and terminal on ulti- dinal range is from near sea level to ca. 1250 m a.s.l.
mate branchlets, ovoid-globose to oblong, 1.52.5 The climate is mild to temperate, mesic or with sum-
12 mm, yellowish green turning orange-brown; mer rainfall in the north and winter rainfall in the
microsporophylls 814, in alternate whorls of 3, pel- south of its range; frost and snow can occur at higher
tate, with denticulate margins, bearing 34 abaxial altitudes.
Conservation branches, linear, 25 mm long on ultimate branch-
lets, 11.5 mm wide, abaxially prominently keeled;
IUCN: LC margins denticulate near slightly broadened obtuse
or acute apex; epistomatic, stomata in two narrow
Uses marginal bands, a few stomata visible near the apex;
abaxial surface weakly verrucose, yellowish green
As a widespread species its timber, being relatively to light green. Pollen cones solitary and terminal
small, was mainly used for fence posts until metal on ultimate branchlets, oblong to cylindrical, 36
replaced wood for this pupose in farming. It has been 1.22 mm, yellowish green turning light brown;
introduced in New Zealand, where it has become microsporophylls 1016, in alternate whorls of 3, pel-
naturalized near Auckland. This introduction had tate, broadly rhombic, with erose-denticulate mar- 247
no horticultural purpose as far as I know. The spe- gins, bearing 34 abaxial pollen sacs near the lower
cies remains relatively rare in botanic gardens and margin. Seed cones terminal on short, thick, leafy
arboreta, although it is available in the UK nursery shoots, mostly aggregated in dense, compact clusters
trade (Grimshaw & Bayton, 2009: 186). on stems and branches, maturing in 11.5 years, per-
sistent, depressed-globose when closed, (10)1218
1220 mm, with concave lower parts of scales,
Callitris roei (Endl.) F. Muell., Syst. Census Austral. smooth, finely rugose when open, whitish grey to
Pl. 1: 109. 1882. Frenela roei Endl., Syn. Conif.: 36. metallic grey, weathering dull grey. Bract-scale com-
1847. Type: Australia: Western Australia, along the plexes in 2 alternate whorls of 3, opening not very
road from Newdegate to Lake King, 35 km E of wide; upper ones largest, (10)1215 610 mm,
Newdegate, A. Farjon 375 (neotype K). Fig. 69, 70 with parallel sides and obtuse apex; smaller ones
wider at base, triangular, with acute apex; a promi-
Etymology nent and acute umbo below the apex of each scale;
abaxial surface concave, becoming rugose, grey;
This species was named after J. S. Roe (17971878), adaxial surface yellowish or more often red-brown
who collected the original specimens, later destroyed with light seed marks. Columella large, triangular or
at W. trilobed, 36 mm long and wide, obtuse. Seeds 36
on each scale, ovoid or triangular, flattened, up to
Vernacular names 10 mm including wings (seed body 35 mm long),
yellowish- or red-brown, with large, whitish con-
Roes Cypress-pine cave hilum; wings 2 on each side, up to 8 4 mm,
unequal in shape and size.
Description
Distribution
Shrubs or small trees 15 m; multistemmed or mono-
podial, branching low, up to 20 cm d.b.h. Bark on SW Western Australia, near the south coast and in
the lower trunk thin, shallowly fissured, hard, slowly parts of the Wheat Belt region.
exfoliating in small flakes, grey. Branches spreading TDWG codes: 50 WAU-WA
or ascending at ca. 45, long and pliant, forming a
broad, rounded, dense or more or less open crown. Ecology
Foliage branchlets rigid and thick, ultimately twisted
and of unequal length, triangular in cross section, In low Eucalyptus woodland (mallee) with Eucalyptus
1.21.5 mm diam., covered with closely appressed spp., other Myrtaceae (e.g. Melaleuca), Proteaceae
leaves, persistent. Leaves in alternate whorls of 3, (e.g. Banksia, Dryandra, Hakea), Leschenaultia sp.,
decurrent, closely appressed, apices appressed or Callitris preissii, and occasionally in open dwarf
some free especially on some leading shoots (whip scrubland (kwongan); on (gravelly) white or grey
shoots), connate but parting with thickening of sand, loam or clay and on laterite (hardpan), also in
disturbed road verges. The altitudinal range is from thick, fissured, stringy, exfoliating in long strips,
50 m to 350 m a.s.l. The climate is characterized by light brown weathering grey. Branches spreading or
warm, dry summers and winter rainfall. ascending, forming an irregular, broad and usually
open crown. Foliage branchlets in tufts at ends of
Conservation main branches, assurgent or erect, ultimate branch-
lets long, apprearing conspicuously articulate with
Due to widespread conversion of the original veg- scale leaves, triangular in cross section, ca. 1.5 mm
etation cover to arable farming this species has diam., covered with closely appressed leaves, green,
suffered a substantial reduction in its area of occu- persistent. Leaves of two types on plants up to 1 m
pancy in the 20th century. Remaining large reserves tall. Adult (scale) leaves in alternate whorls of 3,
248 are mainly concentrated along the south coast, while decurrent, closely appressed, apices appressed or
in the interior these are small and scattered, or have some free especially on some leading shoots (whip
a limited protective status as road verge reserves shoots), connate but parting with thickening of
of semi-natural vegetation. Habitat degradation is branches, linear, 37 mm long on ultimate branch-
mainly caused by increasing salinity of soils in the lets, 0.71 mm wide, obtusely keeled abaxially; mar-
region; along roads fires also have increased, but it is gins erose-denticulate near obtuse or acute apex;
unknown how this species reacts to this. It appears epistomatic, stomata in two narrow marginal bands;
in some situations to react positively to mechanical abaxial surface smooth, green. Pollen cones solitary
soil disturbance along roads (personal observations, and terminal on ultimate branchlets, ovoid or ovoid-
1997) and should be at least mildly tolerant of fire. oblong, 35 1.52.5 mm, yellowish green turning
IUCN: EN [B2ab (iv)] light brown; microsporophylls 915, in alternate
whorls of 3, peltate-triangular, with erose-denticu-
Uses late margins, obtuse or acute, bearing 46 abaxial
pollen sacs near the lower margin. Seed cones ter-
There are no economic uses recorded of this species. minal on short, thin, leafy shoots, solitary, maturing
in 1 year, caducous (or persistent a short time after
seed dispersal), ovoid-globose to subglobose when
Callitris sulcata (Parl.) Schltr., Bot. Jahrb. Syst. 39: closed, 811 710 mm, with concave lower parts of
16. 1906. Frenela sulcata Parl., Index Sem. Hort. scales when open, smooth, finely rugose when open,
Florent. 1862: 23. 1862; Nothocallitris sulcata (Parl.) greenish maturing to dull brown. Bract-scale com-
A. V. Bobrov & Melikyan, Komarovia 4: 85. 2006. plexes in 2 alternate whorls of 3, not opening very
Type: New Caledonia: Grande Terre, Province Sud, wide; upper ones only slightly larger, 710 56 mm,
C. Moore 5 (holotype K). oblong, thick; lower ones 45 mm wide; a promi-
nent and acute umbo below the apex of each scale;
Etymology adaxial surface rugose, red-brown or dark brown
with inconspicuous seed marks. Columella relatively
The species epithet (Latin sulcatus = grooved) refers large, 34 mm long, irregularly angular, 1.52 mm
to the appearance of the branchlets. wide, narrowed at base. Seeds usually 2 at base of
each upper scale, ovoid or triangular, flattened, 45
Vernacular names 3 mm, light brown, with very small hilum; wings 2
on each side, up to 7 2.5 mm, unequal in shape
Sapin de Comboui (French) and size.
Shrubs or small trees to 610(12) m tall; trunk The two species of Callitris in New Caledonia are
monopodial in trees, sometimes forked, branching probably relicts of the Australian connection about
low, up to 40 cm d.b.h. Bark soon scaly, to 1020 mm 80 Ma, and somewhat resemble C. macleayana in
foliage characters. They retain a 4-whorled phyllo- Callitris verrucosa (A. Cunn. ex Endl.) F. Muell.,
taxis with juvenile acicular leaves into semi-mature Rep. Burdekin Exped.: 19. 1860. Frenela verrucosa
or mature plants. In C. sulcata the transition between A. Cunn. ex Endl., Syn. Conif.: 37. 1847; Callitris
leaf types is abrubt, but phyllotaxis changes from 4 preissii Miq. subsp. verrucosa (A. Cunn. ex Endl.) J.
to 3 whorls earlier than leaf type. Cones are only Garden, Contr. New South Wales Natl. Herb. 2 (5):
borne on branchlets with 3-whorled phyllotaxis and 375. 1957; Callitris preissii Miq. var. verrucosa (A.
mature type leaves. Cunn. ex Endl.) Silba, Phytologia Mem. 7: 17. 1984.
Type: Australia: New South Wales, Lachlan River,
Distribution A. Cunningham 372 (holotype K). Pl. 9
plate 9. Callitris verrucosa. 1. Branch with foliage and seed cone. 2. Section of branchlet with leaves.
3. Branchlet with pollen cones. 4. Microsporophyll with pollen sacs. 5. Immature seed cone. 6. Branch with
seed cones. 7, 8. Seeds.
Uses outside a few botanic gardens. There are two foliage
colour forms occurring in the wild, often together:
The wood of this species has similar properties to yellowish green and glaucous green; these could be
that of its congeners, but the shrubby habit makes it selected and enhanced under cultivation.
unsuitable for timber. It is not known in cultivation
252
Calocedrus Kurz, J. Bot. 11: 196. 1873. Type: Calocedrus macrolepis Kurz
(Cupressaceae).
Greek: calo = beautiful, so beautiful cedar; cedrus 1b. Seed cones when mature (4)522 mm long;
is a classical name applied to certain conifers with 1 or 2 seeds developing on each fertile scale 2
fragrant wood, it is not clear which! 2a. Seed cones on long branchlets with numerous
modified monomorphic scale leaves, 1022
Description 47 mm; normally 1 seed developing on each
fertile scale C. macrolepis
Evergreen, monoecious trees with a monopodial 2b. Seed cones on short branchlets with few modi- 253
trunk. Resin cavities in leaves. Bark fissured or fur- fied scale leaves, less than 16 mm long; 1 or 2
rowed, fibrous, stringy, exfoliating in long plates or seeds developing on each fertile scale 3
strips. Branches curved down or spreading to assur- 3a. Leaves and seed cones very small, the latter
gent, forming a pyramidal or sympodial crown. (4)56(7) (2.5)34 mm; normally 2 seeds
Fastigiate forms mostly restricted to trees in culti- developing on each fertile scale C. rupestris
vation. Foliage branches plagiotropic; (sub)ultimate 3b. Leaves and seed cones larger, the latter 1015
branchlets flattened, covered with scale leaves; lat- 46 mm; normally 1 seed developing on each
eral branchlets continually deciduous. Leaves decus- fertile scale C. formosana
sate, appearing in whorls of 4, decurrent, imbricate,
dimorphic, with oblong to obtrullate facials and
slightly broader, conduplicate laterals, very variable Calocedrus decurrens (Torr.) Florin, Taxon 5 (8):
in size dependent on growth of branchlets; margins 192. 1956. Libocedrus decurrens Torr., Smithsonian
entire; apices incurved or recurved, those of lateral Contr. Knowl. 6 (2): 7. t. 3. 1853. Type: USA:
leaves meeting those of facials; stomata mostly on California, Shasta Co., [Upper waters of the
the underside of lateral leaves in two lines or bands, Sacramento River], J. C. Frmont 492 (holotype
glands inconspicuous near apex of facials, absent US?, isotype MO). Fig. 71, 72
on laterals. Pollen cones solitary, terminal, cylindri-
cal; microsporophylls (6)814, decussate, with 35 Etymology
abaxial pollen sacs. Seed cones solitary, terminal,
1035 413 mm, caducous. Bract-scale complexes The species epithet refers to the decurrent leaves on
in 3(4) decussate pairs, the proximal pair small, primary shoots.
recurved, sterile, the middle pair spreading, fertile,
the distal pair fused, sterile. Seeds 12 per fertile Vernacular names
scale, with 2 very unequal wings. Seedlings with 2
cotyledons. Incense Cedar, California Incense Cedar, White
Cedar, Californian Post Cedar, Bastard Cedar;
4 species. Cedro incienso (Spanish)
Distribution Description
W coast of North America, from Oregon to Baja Trees to 60(69) m tall; trunk monopodial, broadly
California; Taiwan, SW China, mainland SE Asia. based or buttressed, up to 34.5 m d.b.h. Bark on
lower trunk deeply furrowed, fibrous, stringy, exfo-
Key to the species of Calocedrus liating into long plates or strips, dark red-brown.
Branches curved down below, then spreading,
1a. Seed cones when mature 2035 mm long; nor- assurgent near the top, long persistent; higher order
mally 2 seeds developing on each fertile cale branches pendulous below, forming a columnar to
C. decurrens pyramidal, eventually domed and more open crown.
Foliage branches numerous, plagiotropic, spreading Ecology
or erect near the top, ultimately flattened, covered
with green leaves, lateral branchlets deciduous in In mixed conifer forest with Pinus jeffreyi, P. pon-
23 years. Leaves decussate, appearing in whorls of derosa, P. lambertiana, P. monticola, Abies con-
4, imbricate; laterals partly covering facials, decur- color, A. grandis, A. magnifica, and Pseudotsuga
rent, (weakly) dimorphic, conduplicate, linear-lan- menziesii, locally with Sequoiadendron giganteum,
ceolate, with slightly incurved acute apex; facials Chamaecyparis lawsoniana, Tsuga heterophylla or
slightly shorter than laterals, oblong to obtrullate, Thuja plicata, and in drier southern sites with Pinus
obtuse-acuminate; leaves persisting several years, coulteri and Pseudotsuga macrocarpa. The under-
variable in size, from 2 1.5 mm on ultimate lateral growth of these mixed conifer forests varies mostly
254 branchlets to 15 3 mm on leading (whip) shoots; with altitude and edaphic conditions and is diverse,
margins entire; amphistomatic, abaxial stomata on especially on ultramafic rocks, with Arctostaphylos
laterals in two narrow bands, more numerous on the patula, A. viscida, Ceanothus cordulatus, C. inte-
underside, stomata on facials restricted to sides and gerrimus, C. parvifolius, Castanopsis sempervirens,
hidden in grooves between leaves; glands weakly Gaultheria shallon and many other shrubby species.
developed, most conspicuous near apex of facials, all In most conifer forest associations C. decurrens is a
exposed faces light green or sometimes dark green. relatively minor component, where it often occu-
Pollen cones solitary and terminal on more or less pies open canopy stands on hot, dry sites. In the
pendulous branchlets, oblong, 68 23 mm, yel- Sierra Nevada Mixed Conifer Forest it may play a
low turning brown; microsporophylls 1014, decus- much greater role in the canopy locally. Other forest
sate, peltate-orbicular with denticulate margins types include also Quercus spp., Castanopsis chryso-
and acutish apex, bearing 34 abaxial pollen sacs phylla, Lithocarpus densiflorus and Arbutus menzie-
near lower margin. Seed cones terminal on more or sii, together with conifers. The altitudinal range of
less pendulous branchlets with mostly unmodified Calocedrus decurrens is from 50 m to 2010 m a.s.l.
leaves (except for 34 subtending pairs of small scale in the north and between 910 m and 2960 m a.s.l. in
leaves), solitary, often numerous, maturing within the south of its range. This species is rather tolerant
1 year, caducous, ovoid-oblong or oblong when to soil types, with a huge range of pH values, but the
closed, (15)2035 813 mm, smooth or striated, soil usually well drained; it is only rare on limestone.
turning red-brown. Bract-scale complexes in 3(4) It tolerates hot, dry summers well, but is equally
decussate pairs; proximal pair undeveloped and insensitive to frost and snow cover.
sterile, with recurved bracts; middle pair spread-
ing wide, fertile, slightly curved, abaxially convex, Conservation
with a minute subapical bract tip and 2 depressed
seed marks near the base of each scale; distal pair IUCN: LC
infertile, straight, laterally flattened and fused to a
flat plate with 2 small seed marks and apical bract Uses
tips; latter two pairs 1830 710 mm, their adaxial
faces rugose, light brown. Seeds oblong, slightly flat- Incense cedar is an important timber tree. The wood
tened, tapering to apex, 812 34 mm, light whitish is used for the manufacture of pencils and in build-
brown, with 2 wings of unequal shape and size; larg- ing for exterior siding and windows of houses and
est wing 1823 mm longwing; smallest a narrow strip garden amenities like fences and trellises. As with
with a free apex. many taxa in this family, the wood is decay resistant,
which makes it especially useful for these purposes
Distribution in the wetter coastal areas of the Pacific States in
the USA. It is also widely planted as an ornamen-
USA: California, Oregon; NW Mexico: Baja tal tree and only then often grows into a fastigiate
California Norte. habit (probably mostly as the cultivar Columnaris
TDWG codes: 73 ORE 76 CAL 79 MXN-BC which was named by Beissner Libocedrus decurrens
var. columnaris but does not occur in nature). hidden margins, on underside facials in two bands;
Incense cedar performs well in urban settings as it glands inconspicuous near apex of facials or absent;
is relatively tolerant of air pollution. In horticulture exposed faces green or sometimes dark green, with
a number of cultivars are known, especially with glaucous green stomatal bands. Pollen cones solitary
variegated or differently coloured foliage. Despite its and terminal on ultimate branchlets, oblong, 45
common name, it is not used for incense burning, 22.5 mm, reddish turning brown; microsporophylls
although its foliage is fragrant. (6)810, decussate, peltate-orbicular with denticu-
late margins and acutish apex, bearing 23 abaxial
pollen sacs near the lower margin. Seed cones ter-
Calocedrus formosana (Florin) Florin, Taxon 5 minal on more or less pendulous branchlets with
(8): 192. 1956. Libocedrus formosana Florin, Svensk ultimately slightly modified (shortened) leaves, soli- 255
Bot. Tidskr. 24: 126. 1930; Calocedrus macrolepis tary, often numerous, maturing within 1 year, cadu-
Kurz var. formosana (Florin) W. C. Cheng & L. K. cous, oblong when closed, 1015 46 mm, smooth
Fu, Fl. Reipubl. Pop. Sin. 7: 327. 1978. Type: Taiwan: or slightly rugose, often glaucous, turning brown.
[Taihoku province (planted)], E. H. Wilson 10960 Bract-scale complexes in 3 decussate pairs; proxi-
(lectotype K). Fig. 73 mal pair undeveloped and sterile, with recurved
bracts; middle pair spreading slightly, fertile, slightly
Etymology curved, abaxially convex, with a small subapical
recurved bract tip and 1 depressed seed mark near
The species epithet indicates Taiwan, formerly base; distal pair infertile, straight, laterally flattened
known as Formosa; see also Latin: formosus = hand- and fused to a flat plate with apical bract tips; latter
some or well formed. two pairs 1013 45 mm, their adaxial faces soft,
smooth, light brown. Seeds 1(2) per cone, ovoid,
Vernacular names flattened, acutish at apex, 45 1.82 mm, brown,
with 2 wings of unequal shape and size; largest wing
Taiwan Incense-cedar; Taiwan cui bai (Chinese) 1012 mm long; smallest wing a narrow strip with a
free apex or virtually absent.
Description
Taxonomic notes
Trees to 2025 m tall; monopodial, up to 23 m
d.b.h. Bark on lower trunk furrowed, fibrous, exfoli- This taxon was treated as a variety of C. macrolepis
ating into long plates or strips, red-brown. Branches in Flora of China 4 (1999); in the Flora of Taiwan (Li,
often few and thick, spreading, ascending near the 1975) C. formosana is recognized as a species. The
top, long persistent, higher order branches spread- morphological differences are small but constant
ing, forming a pyramidal, eventually broad and and C. formosana only occurs in Taiwan.
more or less flat-topped crown. Foliage branches
numerous, plagiotropic, branching alternately, ulti- Distribution
mately flattened and articulate, covered with green
leaves, lateral branchlets deciduous in 23 years. Taiwan: central and northern mountains.
Leaves decussate, appearing in whorls of 4, imbri- TDWG codes: 38 TAI
cate, dimorphic; laterals partly covering facials,
decurrent, conduplicate, linear-lanceolate, with Ecology
recurved-incurved acute apex; facials oblong to
obtrullate, obtuse-acuminate; leaves persisting sev- This species occurs in mixed conifer-broad-leaved
eral years; variable in size, 15 mm long on (sub) climax forest, often as an emergent, associated with
ultimate lateral branchlets to 15 3 mm on leading Pseudotsuga sinensis, Taiwania cryptomerioides,
(whip) shoots; margins entire; amphistomatic, abax- Castanopsis spp., and Cyclobalanus (= Quercus) spp.
ial stomata on laterals in two lines or bands, very few in deep, rich forest soil over shalestone or schist; also
or absent on upperside, much more numerous on in secondary forest and on cliffs and rocky ridges. Its
underside, stomata on upperside facials restricted to altitudinal range is (300)8002000 m a.s.l.
Conservation branches numerous, plagiotropic, branching alter-
nately, ultimately flattened and articulate, covered
This species extends in an area of less than 8000 km2 with green leaves, lateral branchlets deciduous in
and is limited to under ten isolated locations while it 23 years. Leaves decussate, appearing in whorls
suffers a continuing decline in its area of occupancy. of 4, imbricate, dimorphic; laterals partly covering
The causes are timber cutting, transformation of old facials, decurrent, conduplicate, linear-lanceolate,
growth forest to managed production forest, and with recurved-incurved acute apex; facials oblong
expansion of agriculture, especially livestock graz- to obtrullate, obtuse-acuminate; leaf size variable,
ing. Some stands are now protected in reserves and 1.58 mm long on (sub)ultimate lateral branchlets
there is a programme of ex situ conservation to back to 15 3 mm on leading (whip) shoots; margins
256 up the remaining populations. entire; amphistomatic, abaxial stomata on laterals
IUCN: EN [B2ab (ii, iii, v)] in two lines or bands, very few or absent on upper-
side, much more numerous on underside, stomata
Uses on upperside facials restricted to hidden margins, on
underside facials in two bands; glands inconspicu-
The valuable wood of this tree is in demand for con- ous near apex of facials or absent, exposed faces light
struction purposes, mainly at a local or regional level. green with whitish green stomatal bands. Pollen
It is rare in cultivation despite the fact that it can sur- cones solitary and terminal on ultimate branch-
vive winters in e.g. England provided it is planted lets, oblong, 48 23 mm, purplish red turning
in a sheltered spot; a few recently planted trees at light brown; microsporophylls (8)1014, decus-
the Royal Botanic Gardens, Kew have seed cones sate, peltate-orbicular with denticulate margins and
and are doing well. Although known botanically obtuse-acuminate apex, bearing (3)4(5) abaxial
since 1930, it was apparently introduced to cultiva- pollen sacs near the lower margin. Seed cones ter-
tion only recently (Grimshaw & Bayton, 2009: 187). minal on pendulous (5)1020 mm long branchlets
with modified (shortened) monomorphic leaves in
decussate pairs, solitary, often numerous, matur-
Calocedrus macrolepis Kurz, J. Bot. 11: 196., t. 133. ing within 1 year, caducous, oblong when closed,
1873. Libocedrus macrolepis (Kurz) Benth. & Hook. 1022 47 mm, smooth or slightly rugose, often
f., Gen. Pl. 3 (1): 426. 1880. Type: China: Yunnan, glaucous, turning brown. Bract-scale complexes
Daying River, Hotha, D. J. Anderson s.n. in 3(4) decussate pairs; proximal pair undevel-
(holotype K). Fig. 74 oped and sterile, with recurved bracts; middle pair
spreading slightly, fertile, nearly straight, abaxi-
Etymology ally convex, with a small subapical recurved bract
tip and 1(2) depressed seed marks near the base;
The species epithet (Greek/Latin: macro = large; lepis = distal pair infertile, straight, laterally flattened and
scale) refers to the size of the seed cone scales. fused to a flat plate with apical, curved bract tips;
the latter two pairs 1020 46 mm, adaxial faces
Vernacular names soft, smooth, light brown. Seeds 2(4) per cone,
ovoid-oblong, slightly flattened, acute at apex, 56
Chinese Incense-cedar; cui bai (Chinese) 23 mm, brown, with 2 wings of unequal shape and
size; largest wing 1215(20) mm long; smallest wing
Description a narrow strip with a free apex or virtually absent.
Relatively common in montane mixed evergreen Trees to 25 m tall; trunk to 11.2 m d.b.h. Bark 812
conifer-broad-leaved forest dominated by Fagaceae mm thick, fissured on large trunks, fibrous and
and with scattered conifers, e.g. Cunninghamia, exfoliating in longitudinal strips, resinous, greyish
Dacrycarpus, Keteleeria and Pinus; also in rocky brown to brown. Branches spreading and ascending,
places and much planted in roadsides and field forming a broadly rounded crown. Foliage branch-
margins. The altitudinal range is from ca. 800 m to lets spreading and ascending, arranged in a plane,
2000 m a.s.l. covered with scale leaves, flattened, uniformly green
or with very indistinct, whitened stomatal bands on
Conservation the underside. Leaves scale-like, decussate, in whorls 257
of 4, decurrent at base, broadly obtuse to obtuse at
This species has been under threat from cutting for apex, dimorphic; facial pair flattened, (1)26(7)
both timber and firewood, especially in more acces- mm long, (1.5)22.5 mm wide; lateral pair condu-
sible areas in Yunnan. In that province it is still fairly plicate, boat-shaped, (1.5)26(7) mm long, (0.3
common; in its outlying stations, e.g. Hainan Island )0.50.75(1) mm wide, their margins overlapping
and Viet Nam, it is more scarce to rare. Protective with facial leaves, eglandular. Pollen cones terminal,
measures were suggested in the form of reserves as solitary, cylindrical, (4.5)56 mm long, 1.52(2.2)
well as ex situ cultivation in Fu & Jin (eds., 1992). mm wide; microsporophylls decussate, in 811 pairs
IUCN: NT (the lowest 24 pairs sterile), 0.81.2 mm long, 11.2
mm wide, with minutely erose margins, broadly
Uses obtuse or rounded at apex, light green turning light
brown, each with 26 broadly ovoid to subglobose,
In China this species is considered suitable for affor- 0.30.4 mm wide pollen sacs. Seed cone-bearing
estation of deforested lands in its native area because branchlets terete or 4-angled, 0.51.5 mm long, with
of its easy germination and light-demanding proper- 68(12) imbricate, obtuse scales. Seed cones subter-
ties combined with rapid growth. Its wood has good minal, solitary or paired at apex of lateral branch-
properties, e.g. durability, but trees tend to be much lets, ovoid, (4)56(7) mm long, (2.5)34 mm
branched especially when grown in open vegetation. wide, dehiscent when mature (in first year), with 4
It is less hardy than C. decurrens or C. formosana but decussate, flattened scales (very rarely with 2 addi-
is in cultivation in parks and large gardens in China tional, basal scale rudiments). Bract-scale complexes
and elsewhere in SE Asia. somewhat coriaceous, green, becoming woody and
brown, broadly ovate, 46 mm long, 2.54 mm wide;
basal 2 scales fertile, dehiscent, normally 2-seeded,
Calocedrus rupestris Aver., Hiep & L. K. Phan, rarely with 1 seed; apex rounded, incurved, some-
Proc. Nat. Conf. Life Sci. Thai Nguyen Univ. Sept. times with an indistinct, slightly flattened or con-
2004: 41. 2004. Type: Viet Nam: Bac Kan Prov., cave, subapical plate with a rugose surface, rarely
Na Ri District, Na Bo, L. V. Averyanov et al. 5441 with a small central umbo (bract tip); apical pair
(holotype HN). Fig. 75 of scales sterile, connate. Seeds ovoid or sub-ovoid,
acute at apex, slightly flattened, including the two
Etymology relatively large but very unequal wings 45 mm long.
Cathay is an old name for (northern) China. ical, 68 mm long, not resinous; bud scales ovate;
basal scales smaller than the apical scales, light yel-
Description lowish brown, deciduous. Leaves spirally arranged,
on short shoots seemingly in tufts, spreading radi-
See the species description. ally, densely crowded near shoot apex, directed for-
ward, (1.8)2.84.5(5.5) cm long, (2.2)2.53 mm
Distribution wide, more or less petiolate at base, linear, flattened, 259
longitudinally grooved and green above, with 2 glau-
As for the species. cous white bands below; margins slightly recurved,
ciliate in young leaves; apex obtuse. Stomata in two
bands separated by a midrib below, in ca. 15 rows on
Cathaya argyrophylla Chun & Kuang, Acta Bot. each side. Pollen cones lateral, near shoot apex, pen-
Sin. 10 (3): 246. 1962. Pseudotsuga argyrophylla dant, with involucre of light brown, large scales, 36
(Chun & Kuang) Greguss, Bot. Kzlem. 57: 54. cm long, yellowish. Seed cones lateral, at about right
1970; Tsuga argyrophylla (Chun & Kuang) de angles from shoot, short pedunculate, ovoid oblong,
Laub. & Silba, Phytologia Mem. 7: 75. 1984. Type: with acute apex, 35 cm long, 1.52.5 cm wide with
China: Guangxi, Guangfu, [ad regionem sylvati- slightly opened seed scales, greenish, maturing to
cam Kwangfuensem], Kwangfu Lingch Exped 198 orange brown, ripening to chestnut brown. Bracts
(holotype IBSC, isotype PE). Fig. 76, 77 triangular, with a long point, 58 mm long, included.
Seed scales 1316, orbicular or ovate, 1.52.5 cm long
Etymology and 12 cm wide at mid-cone; surface longitudinally
striated or furrowed, initially puberulent, later gla-
The species epithet is derived from the Greek brous; upper margin entire, soon becoming erose;
argyr- = silver and phyla = leaf. base very short pedicellate. Seeds obovoid, with an
acutish base, 56 34 mm, dark olive with pale
Vernacular names green spots; seed wings adnate, obovate cuneate,
1015 mm long and 46 mm wide, light brown.
yin shan (Chinese)
Taxonomic notes
Description
Cathaya argyrophylla is one of the prime examples of
Trees to 20 m tall, d.b.h. to 4060 cm; trunk mono- a relict conifer which had a very much wider distri-
podial, straight, columnar, sometimes forked in the bution in the geologic past. In the Tertiary Cathaya
top; crown pyramidal, often irregular, open. Bark occurred in Europe, Russia, and Canada, and the
grey, becoming scaly, lower down the trunk of older genus is rather common as a fossil in the Miocene
trees breaking into irregular plates. Branches of first lignite (brown coal) deposits of Germany. Its taxo-
order long, heavy, spreading horizontally or slightly nomic position has been the subject of some dispute
ascending; branches of second order slender, branch- in the past, but more recent phylogenetic studies in
ing irregularly. Branchlets slender, firm, yellowish, the Pinaceae have confirmed its status as a genus,
becoming grey in second year; surface with ridges probably related to Pseudotsuga in a pinoid clade.
ending in weak pulvini, divided by grooves, glabrous,
but very young shoots puberulent; leaf scars angular Distribution
circular; no real short shoots (weak dimorphism),
but alternately long and short growth of shoots, the China: Chongqing (Nanchuan Xian, Wulong Xian),
lateral (weaker) shoots forming 13 sequences with NE Guangxi (Jingxiu Yaozu Zizhixian, Longsheng
fewer and shorter leaves. Vegetative buds ovoid con- Gezu Zizhixian), Guizhou (Daozhen Xian, Jiangkou
Xian, Tongzi), S Hunan (Chengdu, Guidong Xian, until very recently. It is a relatively rare conifer, but
Luohandong, Xinning Xian, Zixing Shi). its extent of occurrence (EOO) is now known to
TDWG codes: 36 CHC-CQ CHC-GZ CHS-GX encompass four provinces in south-central China
CHS-HN and herbarium collections (in China) are known
from at least 10 localities. It is usually growing on
Ecology inaccessible slopes and ridges and is not consid-
ered to be a valuable timber resource due to small
On medium high mountains, at elevations between or medium size and poor shape in logging terms.
(900)1200 m and 1900 m a.s.l. The soils are the Several localities are within reserves and enjoy legal
widely distributed red and yellow earths of humid, protection. A re-assessment in 2010 established its
260 warm temperate to subtropical China. Wang (1961) conservation status as Vulnerable on the basis of
has mentioned C. argyrophylla as a rare conifer occur- limited known distribution.
ring in the evergreen sclerophyllous broad-leaved for- IUCN: VU (D1)
est type. This forest type is dominated by numerous
species of Fagaceae with mostly small, ovate lanceo- Uses
late, coriaceous leaves. However, from the altitudinal
range of the species it is likely that it occurs in an eco- This species is in cultivation through several for-
tonal type between the sclerophyllous and deciduous estry institutes and botanic gardens in China; out-
broad-leaved forest types. Other conifers with which side China it is still very rare in collections and no
it occurs are almost certainly Pinus fenzeliana (syn. mature plants exist in these. It has only recently been
P. kwangtungensis), and possibly also Tsuga chinensis freed of its official embargo, which was not based in
and Nothotsuga longibracteata. considerations of conservation of a rare species but
in geopolitics. Lifting of these strictures may have
Conservation begun with the donation of some seedlings by Prof.
Fu Likuo when he visited the Royal Botanic Garden,
After its discovery in the 1950s, for many years this Edinburgh in 1998 for work on Volume 4 of the Flora
monotypic genus was considered to be an extremely of China. Although the species is now available in
rare conifer. Even herbarium specimens were very the trade in both Europe and the USA, it will prob-
few and virtually nothing of it had reached botanic ably remain a tree for collectors gardens only, as it
gardens and institutional herbaria outside China has no remarkable horticultural merit.
Cedrus Trew, Trait Arbr. Arbust. 1: 139. 1755 (nom. cons.). Type: Cedrus libani
A. Rich. (Pinaceae).
Cedrus is the classical Latin name for the (true) junct) regions. If the genus once had a more continu-
cedars; however, it was also in use for (fragrant) ous distribution, with populations bridging the great
wood of other conifers. gaps now extinct, it is evident that this fragmentation
happened long ago. Such separation leads inevitably to
Description genetic differences between the remnant populations,
and eventually to separate species. A comprehensive
Monoecious evergreen trees with a columnar trunk. genetic (DNA based) study of the genus is necessary, 261
Resin canals in bark, leaves, and seed cones, resin but has not yet been undertaken and published.
cysts in wood. Young trees branching at regular inter-
vals, later several branches of the first order become 3 species.
codominant with the main stem, except in C. deo-
dara, where the apical dominance is maintained Distribution
(Massarts model). Bark on large trunks breaking into
small, irregular plates and becoming longitudinally Disjunct: Atlas Mountains of Morocco and Algeria;
fissured. Shoot dimorphism pronounced, with long, Lebanon, Syria, SE Turkey, Cyprus; Himalaya.
slender leading shoots and lateral, short spur shoots;
a terminal bud forming a long shoot, and several lat- Key to the species of Cedrus
eral buds forming short shoots. Leaves linear-acicular,
rigid, usually slightly narrowed near base and taper- 1a. Leading shoot and tips of branches drooping.
ing towards an acute apex, amphistomatic, diamond Leaves on short shoots 24.5 cm long. Pollen
shaped in cross section. Pollen cones solitary and cones usually longer than 5 cm C. deodara
more or less erect from apex of short shoots, large, 1b. Leading shoot and tips of branches erect or
catkin-like; pollen bisaccate; phenology marked by spreading. Leaves on short shoots less than 3
late shedding of pollen (September to November), cm long. Pollen cones shorter than 5 cm 2
related to late development of the female strobilus. 2a. Seed cones when fully mature up to 12 cm long
Seed cones similarly situated, distinctly erect, requir- (but sometimes not exceeding 10 cm). Pollen
ing about 1718 months for full development, becom- cones up to 5 cm long. Indigenous to E
ing ovoid to barrel shaped when full grown. Bracts Mediterranean C. libani
small and hidden, not growing with the seed scales. 2b. Seed cones when fully mature up to 8 cm long.
Seed scales helically arranged on a narrowly coni- Pollen cones up to 4 cm long. Indigenous to W
cal rachis and extremely imbricate, large, flabellate, Mediterranean (Algeria and Morocco)
coriaceous or thin woody, dismembering from the C. atlantica
persistent rachis by abcission at maturity. Seeds fully
covered by a membrane on one side and partly cov-
ered with a small portion at other side, membrane Cedrus atlantica (Endl.) Manetti ex Carrire, Trait
continuing in a very large, broad seed wing. Seedlings Gn. Conif.: 285. 1855. Cedrus libani A. Rich. var.
with (5)810(14) cotyledons. atlantica (Endl.) Hook. f., [Cedars Lebanon] Nat.
Hist. Rev., ser. 2, 2: 15. 1862; Cedrus libani A. Rich.
Taxonomic notes subsp. atlantica (Endl.) Batt. & Trab., Fl. Algrie
Tunisie: 397. 1905. Type not designated. Fig. 78
As will be discussed in more detail below with the
species, the morphology commonly deployed to cir- Etymology
cumscribe and differentiate the constituent species of
the genus Cedrus is on critical examination not very The species epithet means from the Atlantic con-
convincing. The three (or four) species traditionally trasting the species with the cedars that grow in the
recognized occur, however, in vastly separate (dis- eastern Mediterranean.
Vernacular names seed wings broad cuneate, 1825 1217 cm, (light)
brown.
Atlas cedar; Cdre de lAtlas (French)
Taxonomic notes
Description
The taxonomic distinction between Cedrus atlantica
Trees to 3035(40) m tall, but at higher elevations and C. libani is at best inconsistent and, when evalu-
usually lower, d.b.h. to 1.52 m; trunk usually mono- ated critically based on the populations growing
podial, columnar, massive, often forked above the in the wild, probably non-existent. Much has been
middle. Bark on trunk cracked and fissured, rough, made of glaucousness of foliage, but this is based
262 dark grey, breaking into flaking plates revealing red- on trees selected for horticulture; glaucous as well
dish brown bark. Branches of first order massive, as green leaved trees occur in Morocco as well as in
the upper ones ascending, the lower horizontal or Turkey, often in the same populations. Numbers of
bent downward; branches of second order crowded, leaves on short shoots and length of leaves not only
spreading in horizontal or descending planes, vary, the values often cited as distinct between the
ascending near the top of the tree; crown in young two species can be found in both distant areas. The
trees broadly conical, in old trees spreading later- smaller maximum size of seed cones in C. atlan-
ally, becoming flat topped. Branchlets short, firm, tica may be correct, but to emphasize this hides the
except leading shoot, which is longer and slender, substantial overlap in cone sizes actually observed
grey green or grey brown, soon turning grey and in natural populations. Crown shape is entirely a
flaking, densely blackish pubescent at first; pulvini question of age and of growing conditions; roughly
on long shoots small, prominent; short shoots thick, speaking: the older the tree the flatter the crown in
scaly, of variable length with age (0.53 cm), assur- both species. In Turkey, trees with pyramidal crowns
gent or erect. Vegetative buds ovoid globular, 23 are known as C. libani subsp. or var. stenocoma,
1.52 mm, not resinous; bud scales broadly ovate, but that distinction is taxonomically doubtful, too.
red brown, dark brown or blackish at apex, decidu- Should we therefore unite the Mediterranean Cedars
ous. Leaves on long shoots spirally arranged, remote, under a single species, C. libani? Should we recog-
near base of long shoot more crowded, radially nize subspecies or varieties? The only true distinc-
spreading, falling in 2nd or 3rd year; on short shoots tion is the great geographical separation, which rules
densely crowded in false whorls, 2045, spreading out gene flow between C. atlantica and C. libani in
radially, (1)1.52.5(3) cm long, 11.5 mm wide, nar- nature. They may become truly distinct species in
rowly linear, straight or curved, diamond shaped in the future.
cross section; apex acute or acuminate; stomata on
all sides, more numerous on two adjacent sides; leaf Distribution
colour glossy dark green or glaucous. Pollen cones
terminal on short shoots, erect, subtended by leaves, NW Africa: Algeria, Morocco (Atlas Mts.).
numerous, soon falling after shedding pollen, 34 TDWG codes: 20 ALG MOR-MO
cm long, straight, later curved, first rose yellow, later
pale brown. Seed cones terminal on short shoots, Ecology
erect, sessile, becoming woody in 2nd year, ovoid or
barrel shaped; apex obtuse or retuse, 58 35 cm, The Atlas cedar occurs on the high maritime ranges
light green, maturing to pale green, with purplish of the Atlas Mountains, at elevations between 1370 m
edges of seed scales, ripening to light (purplish) and 2500 m a.s.l., especially on N and NW exposed
brown; cone rachis persistent, narrowly conical. slopes receiving 1000 to 2000 mm precipitation
Seed scales broad flabellate, thin, coriaceous, length annually, mostly during the winter. At the high
width 23 2.53.5 cm; surface smooth, orange- ridges much snow accumulates. The summers are
brown pubescent at base, glabrous on exposed parts; warm and dry. The soil is usually rocky, calcareous
upper margin entire, slightly incurved; base pedicel- and well drained. The species grows in pure stands,
late. Seeds ovoid conical, 813 46 mm, brown; reaching tree limit on high, exposed ridges, or is
locally mixed with Abies numidica, Taxus baccata, Vernacular names
Quercus faginea, Acer obtusatum, Populus tremula,
Sorbus aria, and S. torminalis (Algeria) or Abies Deodar cedar, Himalayan cedar; deodar (India);
pinsapo var. marocana, Juniperus communis, Taxus xuesong (Chinese)
baccata, Populus tremula, Quercus ilex, and Acer
granatense locally in Morocco. Description
plate 10. Cedrus deodara. 1. Habit of tree. 2. Branch with foliage. 3. Seed cone. 4. Pollen cone and leaves.
5. Leaf. 6. Seed.
brown pubescent at base; upper margin entire, thin, Conservation
slightly incurved; base pedicellate. Seeds ovoid coni-
cal, 1015 57 mm, brown; seed wings broad cune- IUCN: LC
ate, 2030 1521 mm, light brown.
Uses
Taxonomic notes
Himalayan cedar is a very important timber tree in
Reducing Cedrus deodara to a subspecies of C. libani, Pakistan, Kashmir and NW India. Its strong and dura-
as was proposed by Peter Sell (op. cit.), would presum- ble wood is mostly used for construction. Other uses
ably reduce Cedrus Trew to a monotypic species, with are general carpentry and furniture. A fragrant oil
only a few varieties or subspecies recognized under can be distilled from wood chips and sawdust. It was 265
that species. Based on morphology, there is merit in first introduced in England in 1822 as an ornamental
this view, but it may not be the entire truth about tree, but as a park tree it remains less common than
what in this case actually constitutes a distinct spe- the two Mediterranean species. It is generally more
cies. Unlike between C. atlantica and C. libani, there susceptibe to late frost than these and also requires
are consistent, albeit small, morphological differences more moisture. Young trees especially have a distinct
between these two species and C. deodara. More habit with drooping leaders and are commonly used
research into genetic aspects of the problem is needed in gardens. This use has led to the selection of several
before such a far reaching conclusion can be drawn. cultivars, with different habit including dwarf forms
and/or with varying foliage colours; most of these are
Distribution in cultivation in Central Europe.
Greek: kephalos = head; referring to the structure Key to the species of Cephalotaxus
(cone) onto which the seed is attached; Taxus is the
classical Latin name for yews. The number of lines of stomata in a stomatal band
is not necessarily correlated with the width of that
268 Description band, so a magnifying lens 1020 is required to
observe this character. The disposition of leaves (in
Dioecious evergreen shrubs or small trees. Resin a horizontal plane or in a V-formation) can vary
canals (1) in the leaves only. Bark thin, smooth, between fertile and infertile shoots and between
exfoliating in narrow strips. Branches spreading sun-exposed and shaded shoots and is by itself not a
and plagiotropic or ascending, foliage branches reliable character.
terminating in conical, scaly buds. Leaves spirally
inserted, spreading in all directions on erect shoots, 1a. Leaves convex, touching each other closely,
pectinately arranged on lateral, plagiotropic shoots cordate or truncate at base; apex minutely cus-
by alternately twisting of petiolate leaf bases, becom- pidate or mucronate C. oliveri
ing subopposite, flattened, linear-lanceolate, with 1b. Leaves more or less flat, separately disposed,
two prominent stomatal bands separated by a raised cuneate to nearly truncate or twisted petiolate
midrib on the abaxial side. Pollen cones aggregated at base; apex acuminate or cuspidate, some-
in pendulous, more or less globose, short-stalked times mucronate 2
clusters (capitulae) in the axils of leaves, forming 2a. Leaves mostly 3.510(12.5) cm long, curved
opposite or subopposite rows on the underside of lat- upward and outward or downward, forming a
eral foliage branchlets; each small cone consiting of V-formation. Peduncle of seed-bearing struc-
a short rachis with up to 15 peltate microsporophylls ture variable in length, but up to 2025 mm
in spiral arrangement, each with 24 pollen sacs long 3
containing spherical pollen. Seed cones in clustered 2b. Leaves mostly 25(7) cm long, spreading hori-
pairs or groups from axillary buds situated distally zontally or forming a V-formation on shoots
from lateral shoots, (long) pedunculate, consisting with pollen cones. Peduncle of seed bearing
of two sterile and several decussate fertile bracts, structure up to 10(15) mm long 4
the fertile bracts with two axillary, erect ovules, 3a. Leaves (1.5)35 mm wide. Seed-bearing struc-
somewhat swollen at pollination and at that time tures 36 together; seeds including the striated
still lacking an arillus, developing much later. Seeds or ridged aril 1425 915 mm C. fortunei
from 12 fertilized ovules per cone, large, obovoid 3b. Leaves 47 mm wide. Seed-bearing structures
to ellipsoid, surrounded by a fleshy, green aril rip- solitary; seeds including the indistinctly stri-
ening from green to yellowish, reddish or purplish ated aril 3545 2025 mm C. lanceolata
brown. Seed proper large, with a hard, sclerified 4a. Stomatal bands with 1015 lines of stomata; leaf
seed coat. bases cuneate 5
4b. Stomatal bands with 2025 lines of stomata;
8 species. leaf bases obtuse to nearly truncate 6
5a. Pollen cones 67 per capitulum, to 3 mm diam.
Distribution Seeds including the aril ellipsoid, with 6 longi-
tudinal ridges C. sinensis
NE India (Arunachal Pradesh, Assam); Myanmar 5b. Pollen cones 612 per capitulum, 34 mm diam.
[Burma]; China; Korea; Japan; Taiwan; Lao PDR; Seeds including the aril obovoid, smooth or
Thailand; Viet Nam. striated 7
6a. Leaves spreading at 4580 to shoot axis, linear dered by green, slightly revolute leaf margins. Pollen
and abruptly narrowing to a mucronate apex cones situated in rows of capitula on the underside
C. hainanensis of lateral foliage branchlets; each capitulum on a 37
6b. Leaves spreading at 7090 to shoot axis, wid- mm long, scaly peduncle inserted in the upper axil
est below the middle and gradually tapering to of a leaf, subtended by ovate, incurved bracts with
a cuspidate apex C. mannii entire or erose, more or less hyaline margins, bear-
7a. Leaves on shoots with pollen cones forming a ing 614 small, sessile or short pedunculate, globose
V-formation. Peduncle of seed-bearing struc- cones 34 mm diam. Microsporophylls 616 per
ture 515 mm long C. harringtonii cone, each with 34 globose, cream coloured pollen
7b. Leaves on shoots with pollen cones (as other sacs. Seed cones borne 36 together at base of lateral
leaves) spreading more or less horizontally foliage branchlets, on 525 mm long, slender pedun- 269
Peduncle of seed-bearing structure 25 mm cles. Bracts several per cone, reduced, 11.5 mm
long C. latifolia long; fertilized ovules surrounded by an ellipsoid,
green or yellow aril, enclosing the ripening seed,
becoming 1425 915 mm, turning soft, purple and
Cephalotaxus fortunei Hook., Curtiss Bot. Mag. 76: indistinctly striated or longitudinally ridged, with a
t. 4499. 1850. short mucronate apex. Seeds ellipsoid or sometimes
globose, 1324 714 mm.
Etymology
Distribution
This species was named after Robert Fortune, who
introduced it to England in 1848. SW, Central and SE China; N Myanmar [Burma].
TDWG codes: 36 CHC-CQ CHC-GZ CHC-HU
Vernacular names CHC-SC CHC-YN CHS-AH CHS-FJ CHS-GD CHS-GX
CHS-HE CHS-HK CHS-HN CHS-JS CHS-JX CHS-ZJ
Fortunes plum yew; san jian shan (Chinese) 41 MYA
Description Ecology
Shrubs or trees 120 m tall; trunk to 40 cm d.b.h. or Cephalotaxus fortunei occurs as an understorey tree
more, often multi-stemmed. Bark thin, exfoliating in or shrub in mixed broad-leaved (angiosperm) for-
strips, reddish brown. Branches spreading or ascend- ests, in mixed conifer-broad-leaved forests and in
ing, forming a broad crown in trees; shrubs often as coniferous forests (Abies and Picea). It is also com-
wide or wider than tall. Foliage branchlets slender, mon as a shrub or small tree in open thickets and
terete, glabrous, finely grooved between decurrent on roadsides in secondary vegetation. Its altitudi-
leaf bases, green turning yellowish to light brown. nal range is great and extends from 200 m to 3700
Leaves mostly spreading in two rows in a semi- m a.s.l., with C. fortunei var. alpina at the higher
pectinate arrangement, diverging at various angles range between (1100)1800 m and 3700 m a.s.l. In
between 30110 to shoot axis, commonly curved the Lower Yangtze Valley var. fortunei occurs in
upward and down towards apex; (1.5)3.510(12.5) remnants of mixed mesophytic forest with Acer
cm long, linear-lanceolate, straight or slightly fal- spp., Catalpa ovata, Fraxinus chinensis, Ilex latifo-
cate, (1.5)35 mm wide, with a short, twisted petio- lia, Liquidambar formosana, Nyssa sinensis, Quercus
late base, gradually tapering to an acute to cuspidate spp., and many other angiosperm trees. Most of
apex, coriaceous but flexible, dark dull green above, these forest remnants are disturbed and/or replaced
with two whitish green bands below. Midrib on the by secondary shrubby vegetation, in which C. fortu-
adaxial (upper) side in a shallow depression, 0.5 mm nei may recur. In the evergreen broad-leaved forests
wide, obtusely raised, continuous from base to apex, of Fujian, which are also remnants, C. fortunei var.
obtusely raised and continuous on the abaxial side. fortunei occurs in the understorey of oaks (Quercus
Stomata in two broad bands of 1724 intermittent spp.) but mixed with many other tree species, some
white lines separated by a green midrib and bor- of which are conifers like Nageia nagi, Keteleeria
fortunei and Fokienia hodginsii. In S Gansu, Shaanxi TDWG codes: 36 CHC-CQ CHC-GZ CHC-HU
and Sichuan var. alpina occurs in very different for- CHC-SC CHC-YN CHS-AH CHS-FJ CHS-GD CHS-GX
ests dominated by Abies, Picea or Larix, or a mixture CHS-HE CHS-HN CHS-JS CHS-JX CHS-ZJ 41 MYA
of these conifers, between 1800 m and 3600 m a.s.l.
Here it is a subcanopy tree or shrub, often accompa- Conservation
nied by Taxus chinensis and shrubs like Eurya and
Rhododendron. IUCN: LC
The species epithet derives from Hainan, the island In Flora of China 4: 87 (1999), this species is treated
(and now province) of China where this species is as a synonym of Cephalotaxus mannii. However,
native and perhaps indigenous. in a note the co-author Robert Mill considered the
plants from Hainan to be separable as the species 271
Vernacular names named and described by H. L. Li, i.e. C. hainanensis.
Mill gives a statement of morphological differences
Hainan plum yew; hai nan cu fei (Chinese, includes between the two species. Under this narrower spe-
C. mannii in Flora of China 4, 1999). cies circumscription it is possible that other speci-
mens from southern China and northern Viet Nam
Description would have to be included under C. hainanensis,
with consequences for its distribution and hence
Trees to 20 m tall; trunk to 70 cm d.b.h. Bark thin, conservation status. Ying et al. (2004) map this spe-
exfoliating in small or large flakes and strips, light cies as occurring in W Guangdong, Hainan, SE and
brown to reddish brown, weathering grey. Branches W Guanxi, W Yunnan, and even SE Xizang [Tibet].
spreading or ascending, forming a rounded or nar- The populations of Yunnan and Tibet are here
row crown. Foliage branchlets up to 15 cm long, treated as belonging to C. mannii. A critical modern
slender, grooved between decurrent leaf bases, green revision of the genus to resolve these issues is lacking
turning orange-brown. Leaves arranged in two pec- at present.
tinate rows, spreading horizontally and at 4580 to
shoot axis, subopposite, (1.5)24 cm long, slightly Distribution
falcate, 2.54 mm wide, flat, base very short petio-
late or sessile, broadly obtuse to nearly truncate, leaf China: Hainan Island (Jianfeng Ling, Limu Ling,
blade linear and abruptly narrowing to a mucronate Wuzhi Shan, Dawang Ling, Diao Luo Shan).
apex. Midrib thin but prominent and continuous TDWG codes: 36 CHH
from base to apex on the adaxial (upper) side, 0.3
mm wide, conspicuous but more flattened on the Ecology
abaxial side; leaf colour dark green or olive-green
above, two white or glaucous white bands below. Cephalotaxus hainanensis occurs in mixed warm
Stomata in two broad bands of ca. 20 intermittent, temperate to subtropical rainforests in mountainous
white lines, separated by the green midrib and bor- areas of the island of Hainan, where it attains tree
dered by slightly revolute leaf margins. Pollen cones habit and size (1020 m). Nothing has been recorded
situated in rows of capitula on the underside of lat- in non-Chinese literature about associated species
eral foliage branchlets; each capitulum on a (1)35 at present, presumably these forests are diverse. It
mm long, scaly peduncle inserted in the upper axil occurs from sea level to about 1700 m a.s.l. and can
of a leaf, subtended by ovate, incurved bracts with be the dominant tree in some places.
entire or erose, more or less hyaline margins, bear-
ing 68 small, sessile, globose, cream or light yellow Conservation
cones up to 4 mm diam. Microsporophylls 612 per
cone, each with 34 globose, cream coloured pol- Under the narrower circumscription accepted here
len sacs. Seed cones solitary at base of lateral foliage this species is probably restricted to Hainan but its
branchlets, on 410 mm long peduncles. Bracts few entire range remains uncertain. It is threatened by
per cone, reduced, 11.5 mm long; fertilized ovules timber harvesting and especially stripping of bark.
surrounded by an ellipsoid, green aril, enclosing the IUCN: EN [B2ab (iii)]
Uses capitula on the underside of lateral foliage branch-
lets; each capitulum on a 37 mm long, scaly pedun-
The bark of this and other species is stripped to cle inserted in the upper axil of a leaf, subtended by
be used for medicinal purposes in China. It is not ovate, incurved bracts with entire or erose, more or
known to be in cultivation outside a few botanic less hyaline margins, bearing 612 small, sessile or
gardens. short pedunculate, globose cones 34 mm diam.
Microsporophylls 515 per cone, each with 23 glo-
bose, cream coloured pollen sacs. Seed cones borne
Cephalotaxus harringtonii (Knight ex J. Forbes) K. 36 together at base of lateral foliage branchlets, on
Koch, Dendrol. 2 (2): 102. 1873. 515 mm long, slender peduncles with small scale
272 leaves. Bracts several per cone, reduced, 11.5 mm
Etymology long; fertilized ovules surrounded by an obovoid,
green or purplish aril, enclosing the ripening seed,
This species commemorates the Earl of Harrington, becoming 1520(25) 1218 mm, turning soft,
who gave a specimen to the Duke of Bedford, from orange-red to purple and smooth or striated, with
whose pinetum at Woburn Abbey James Forbes a short mucronate apex. Seeds (ob)ovoid to subglo-
described and named it. bose, 1218 812 mm.
Harringtons plum yew, Japanese plum yew; inu- This species has long been known as Cephalotaxus
gaya (Japanese); Picha-nam (Korean) drupacea Siebold & Zucc. and it was introduced to
cultivation in the Netherlands by Von Siebold under
Description that name. However, this name was only validly pub-
lished in 1846, by which time the same species (intro-
Shrubs to small trees 0.510 m tall; trunk d.b.h. of duced by Von Siebold as stated) had been named and
monopodial trees to 40 cm. Habit prostrate to erect published as Taxus harringtonii [harringtonia] by
and spreading; trees with a wide, open and rounded James Forbes in 1839, based in part on a manuscript
crown. Bark thin, exfoliating in narrow strips, grey- received from the nurseryman Joseph Knight. It
brown. Branches spreading, drooping or ascending, was later transferred to Cephalotaxus by Karl Koch.
or prostrate in one variety; foliage branchlets short, Cephalotaxus koreana Nakai was described as a 11.5
stout or slender, glabrous, grooved between decur- m tall, caespitose but non-layering shrub with red
rent leaf bases, green turning orange-brown. Leaves [and] most delicious palatable fruits. Those of C. dru-
more or less disposed in two ranks, often upright pacea (= C. harringtonii) were said to be bitter with a
forming a V-formation especially on branchlets disagreeable smell. Otherwise the two taxa are simi-
with male cones, but variable and not limited to lar and the assertion about edibility apparently being
this species, straight or curved down towards apex, based on the experience and by its nature subjective
(1)2.54(5) cm long, with a short, twisted petiole opinion of one observer is here not considered to be
and cuneate base, linear-lanceolate, 2.54 mm wide, a valid taxonomic character. Cephalotaxus koreana
gradually or more abruptly tapering to an acumi- was said by its author to occur in Korea as well as in
nate or cuspidate apex, coriaceous and more or less Japan, where C. harringtonii can be a shrub as well as a
rigid, green above, with two pale green bands below. small tree. Probably the sugar content of the aril varies
Midrib on the adaxial (upper) side prominent, 0.5 among individuals or (sub)populations of this species.
mm wide, obtusely raised and continuous from base
to apex, on the abaxial side nearly flat, 0.50.7 mm Distribution
wide and continuous. Stomata in two broad bands
on the abaxial side, each with 1015 intermittent North and South Korea, Japan, Taiwan.
white lines, separated by the midrib and bordered TDWG codes: 38 JAP-HN JAP-KY JAP-SH KOR-NK
by flat leaf margins. Pollen cones situated in rows of KOR-SK TAI
Ecology especially the smaller var. nana, which spreads read-
ily by suckers.
Cephalotaxus harringtonii in its tree form is a com-
ponent of both broad-leaved (angiosperm) forest 3 varieties are recognized:
and coniferous forest, or mixed forest, occurring
in the understorey. Cephalotaxus harringtonii var.
nana is a spreading shrub on seaside cliffs as well Cephalotaxus harringtonii (Knight ex J. Forbes)
as in mountains over rocky terrain; var. harringtonii K. Koch var. harringtonii. Taxus harringtonii
can also occur as an upright shrub in dense thickets. Knight ex J. Forbes, Pinetum Woburn.: 217, t. 63.
Only var. wilsoniana in Taiwan is invariably a small 1839, [harringtonia]; Cephalotaxus drupacea
tree in mixed montane forests. It appears that tree Siebold & Zucc. var. harringtonii (Knight ex 273
forms of this species are increasingly common in J. Forbes) Pilg., in Engler, Pflanzenr. IV.5 [18]:
more southern regions where the winters are milder. 102. 1903. Type: Illustration in J. Forbes, Pinetum
The altitudinal range of var. harringtonii is not well Woburnense, t. 66. 1839 (holotype). Fig. 83, 84
documented; one herbarium collection was made at
600 m a.s.l. Variety nana occurs from 10 m near the Cephalotaxus drupacea Siebold & Zucc., Abh. Math.-
coast to 1900 m a.s.l. in the mountains. In Taiwan, Phys. Cl. Knigl. Bayer. Akad. Wiss. 4 (3): 234. 1846.
var. wilsoniana is recorded from 1800 m to 2700 m Cephalotaxus koreana Nakai, Bot. Mag. (Tokyo) 44:
a.s.l., so this is a high montane forest tree. In Japan, 508. 1930.
var. harringtonii is common in the undergrowth of
forests dominated by Abies sachalinensis and in open Description
moorland in colder, northern regions. In S Japan
var. harringtonii occurs in mixed forests with Acer, Shrub to small tree 10 m tall. Leaves often in an
Quercus, Tsuga, Chamaecyparis, Abies, and some- upright V-formation, especially on shoots with pol-
times, also as a smaller subcanopy tree, Podocarpus len cones, (2.5)34.5(5) cm long, 34 mm wide,
macrophyllus. gradually or sometimes more abruptly tapering to
an acuminate or cuspidate apex.
Uses
Distribution
In Japan, the wood of Harringtons plum yew is of
minor commercial importance and traditionally North and South Korea; Japan.
used for tool handles and household utensils. The TDWG codes: 38 JAP-HN JAP-KY JAP-SH KOR-NK
seed arils contain oil which was formerly pressed KOR-SK
from them and used in lamps. This species has long
been cultivated for gardens in Japan, where two of Conservation
the three varieties here recognized are native. It was
sent to the Leiden Botanical Garden by Philipp von IUCN: LC
Siebold in 1829 as C. drupacea and soon distributed
from there by cuttings, arriving in Ghent, Belgium
in 1830. The type of the species C. harringtonii is Cephalotaxus harringtonii (Knight ex J. Forbes)
(was) a plant cultivated in the gardens of the Duke K. Koch var. nana (Nakai) Rehd., J. Arnold Arbor.
of Bedford at Woburn Abbey, Bedfordshire, England 22: 571. 1941. Cephalotaxus nana Nakai, Bot. Mag.
and represented by plate 66 in the book Pinetum (Tokyo) 33: 193. 1919. Type not designated.
Woburnense (Forbes, 1839). It was obtained from
Japan by the nursery of Knight & Perry, Chelsea, Description
London, most likely from seeds obtained from plants
in cultivation. It has now been spread throughout the Shrubs with upright, suckering stems and layer-
northern hemisphere and beyond and is quite com- ing lower branches; leaves more or less horizon-
monly seen in gardens and parks, including its var. tally arranged, 13(3.5) cm long, 23.5 mm wide,
nana and several cultivars. It is suitable for hedges, abruptly cuspidate.
Distribution Distribution
Conservation Ecology
This species was named after George Mann, who Taxonomic notes 277
collected the type specimen.
This species has sometimes been treated in a wider
Vernacular names sense to include C. hainanensis, e.g. in Flora of
China 4 (1999), but this taxon is here treated as a
Manns plum yew; hai nan cu fei (Chinese), inh distinct species. Cephalotaxus is a difficult genus
tng, Phi ba mi (Vietnamese) morphologically and a comprehensive critical revi-
sion including DNA-based analyses is long overdue.
Description
Distribution
Trees to 30(50?) m tall; trunk to 70(120) cm d.b.h.
Bark thin, exfoliating in small or large flakes and China: Guangdong (Xingyi), Guangxi, Hainan, SE
strips, light brown to reddish brown, weathering Yunnan, SE Xizang [Tibet]; N Myanmar [Burma];
grey. Branches spreading or ascending, forming a NE India: Arunachal Pradesh, Megalaya (Khasi
rounded or narrow crown. Foliage branchlets up to Hills, Jaintia Hills), Assam (Manipur, Nagaland);
25 cm long, slender, grooved between decurrent leaf NW Thailand; Viet Nam; Lao PDR.
bases, green turning orange-brown. Leaves arranged TDWG codes: 36 CHC-YN CHH CHS-GD CHS-GX
in two pectinate rows, spreading horizontally and CHT 40 ASS-MA ASS-ME ASS-MI ASS-NA EHM-AP
at 7090 to shoot axis, subopposite, (1.5)2.55( 41 LAO MYA THA VIE
6) cm long, straight or commonly slightly falcate,
(2)2.54 mm wide, flat, base very short petiolate Ecology
or sessile, broadly obtuse to nearly truncate, leaf
blade widest below the middle and gradually taper- This widespread species occurs in mixed evergreen
ing to a cuspidate apex. Midrib thin but prominent or deciduous forests, often in ravines. Its altitudinal
and continuous from base to apex on the adaxial range is from 500 m to 2000 m a.s.l. In China trees
(upper) side, 0.4 mm wide, conspicuous but more are described as not reaching taller than 20 m (Flora
flattened on the abaxial side; leaf colour dark green of China 4: 87, 1999), but in Thailand trees up to 50
or olive-green above, two white or glaucous white m tall have been reported. That maximum figure is
bands below. Stomata in two broad bands of ca. certainly in need of verification, but trees to 30 m
2025 intermittent, white lines, separated by the tall commonly occur in the undisturbed evergreen
green midrib and bordered by slightly revolute leaf submontane rainforests of Thailand and Viet Nam.
margins. Pollen cones situated in rows of capitula Cephalotaxus mannii can occur on both silicate rocks
on the underside of lateral foliage branchlets; each and limestone. It is often associated with Nageia wal-
capitulum on a (1)35 mm long, scaly peduncle lichiana, Taxus wallichiana, Dacrycarpus imbricatus
inserted in the upper axil of a leaf, subtended by and Podocarpus neriifolius on soils derived from the
ovate, incurved bracts with entire or erose, more or silicate rocks, and with Pseudotsuga sinensis, Nageia
less hyaline margins, bearing 68 small, sessile, glo- fleuryi, Pinus kwangtungensis, Podocarpus pilgeri,
bose, cream or light yellow cones up to 4 mm diam. Taxus chinensis, Fokienia hodginsii, and Amentotaxus
Microsporophylls 612 per cone, each with 34 spp. on karst limestone in N Viet Nam and S China.
In both types of habitat angiosperms play an impor- Description
tant role with numerous tree and shrub species as
well as epiphytes. Shrubs or small trees to 4 m tall. Bark thin, becom-
ing scaly with thin flakes, yellow weathering greyish
Conservation brown. Branches spreading to ascending, forming
a bushy crown. Foliage branchlets slender, lateral
Globally this species is considered Vulnerable; in branchlets (sub)opposite, often in a plane, up to 15
China it is considered Endangered, due to the range- cm long but usually shorter than 10 cm, grooved
wide reduction of forests. Throughout its range, C. between decurrent leaf bases, yellowish green turn-
mannii is restricted to small populations in which ing yellow to light brown. Leaves in two opposite,
278 the largest trees (to 30 m tall and over 100 cm d.b.h.) distichous ranks at 6080 to shoot axis, mostly of
are often targeted for their timber. Stripping of bark equal length and touching each other from base
is often fatal to the trees and this type of harvest, like to near apex, (1.5)23(3.5) cm long, (2.3)2.7
logging due to slow growth, is unsustainable. The 3.5(4) mm wide, straight or slightly falcate and con-
principal threat, however, is conversion of habitat to vex, sessile or nearly so, base truncate or weakly cor-
agriculture and resulting severe forest fragmentation. date, flat margins parallel to near the short, minutely
There are not enough forest reserves of sufficient size cuspidate or mucronate apex, leaf texture coria-
and integrity to safeguard this species at present (Fu & ceous, slightly rugose, stiff; leaf colour dull green
Jin, 1992; Nguyen Tien Hiep et al., 2004). above, pale green below with two whitish bands.
IUCN: VU (A2cd) Midrib on the adaxial (upper) side narrow (less
than 0.4 mm wide), raised but fading towards apex,
Uses on the abaxial side 0.70.8 mm wide, flat. Stomata
in two broad bands of 1317 intermittent lines sepa-
This conifer produces high quality insect resistant rated by the abaxial midrib and two 0.7 mm wide
timber which is used for quality furniture, fine crafts margins and of equal width to these. Pollen cones
and tool handles. The seeds have medicinal qualities situated in rows of capitula on the underside of lat-
and in Hainan the bark is used to treat fever. The eral foliage branchlets; each capitulum on a 13 mm
species is eminently suitable for use in horticulture, long, scaly peduncle inserted in the upper axil of a
but is seldom seen in cultivation outside Asia. Some leaf, subtended by ovate, incurved bracts with entire
plants may be grown under its taxonomic synonym, or erose, more or less hyaline margins, bearing 36
C. griffithii Hook. f., which was originally described small, sessile, globose, pink or light brown cones up
from the Mishmi Hills in Assam, India. At the Royal to 2 mm diam. Microsporophylls 49 per cone, each
Botanic Garden, Edinburgh cuttings are being prop- with 3(4) globose, pink coloured pollen sacs. Seed
agated from sources in Viet Nam. cones solitary or in groups at base of lateral foli-
age branchlets, on 37 mm long, curved peduncles.
Cephalotaxus oliveri Mast., Bull. Herb. Boiss. 6: Bracts several per cone, reduced, 11.5 mm long;
270. 1898. Type: China: Hubei, Changyang T. Z., A. fertilized ovules surrounded by an obovoid, green
Henry 7479 (lectotype K, designated here). or grayish aril, enclosing the ripening seed, becom-
ing 2227 1418 mm, turning soft, orange-red to
Etymology red with longitudinal striation. Seeds (ob)ovoid to
ovoid, sometimes subglobose, 1823 1015 mm,
The species epithet commemorates F. W. Oliver, a with a mucronate apex.
botanist who collected in China.
Taxonomic notes
Vernacular names
Records of this species from Viet Nam are referable
Olivers plum yew; bi zi san jian shan (Chinese) to C. mannii.
Distribution Cephalotaxus sinensis (Rehd. & E. H. Wilson)
H. L. Li, Lloydia 16 (3): 162. 1953. Cephalotaxus
China: Chongqing, N Guangdong, Guizhou, W drupacea Siebold & Zucc. var. sinensis Rehd. &
Hubei, Hunan, E Jiangxi, S and W Sichuan (espe- E. H. Wilson, in Sargent, Pl. Wilson. 2: 3. 1914;
cially Emei Shan), E Yunnan. Cephalotaxus harringtonii (Knight ex J. Forbes)
TDWG codes: 36 CHC-CQ CHC-GZ CHC-HU K. Koch var. sinensis (Rehd. & E. H. Wilson)
CHC-SC CHC-YN CHS-GD CHS-HN CHS-JX Rehd., J. Arnold Arbor. 22: 571. 1941. Type: China:
W Sichuan, Baoxing, [Mupin], E. H. Wilson 1115
Ecology (holotype A).
Retinispora Siebold & Zucc., Fl. Japon. 2 (5): 36. Key to the species of Chamaecyparis
1844. Type: Retinispora obtusa Siebold & Zucc.
[Chamaecyparis obtusa (Siebold & Zucc.) Endl.] 1a. Seed cone scales 48(10) in number, mature
cones 48 mm diam 2
Greek: chamae- = lowly, creeping; cyparis = cypress. 1b. Seed cone scales 814(16) in number, mature
cones 714 mm diam 3 281
Description 2a. Smallest branchlets plagiotropic (in flat sprays);
ultimate branchlets often unilateral C. pisifera
Evergreen, monoecious trees; trunk monopodial. 2b. Smallets branchlets irregularly disposed; ulti-
Resin cavities in leaves. Bark fissured, fibrous or mate branchlets alternating C. thyoides
scaly, exfoliating in long strips or flakes, reddish 3a. Seed cones widely open, with gaps between
brown. Branches slender, persistent, forming a pyra- scales as wide as or wider than scales. Leaf
midal, conical or rounded crown (Massarts model). gland present; leaf apices of laterals more or less
Fastigiate forms restricted to cultivation. Foliage acute C. lawsoniana
branches plagiotropic, drooping or pendulous, (sub) 3b. Seed cones not widely open, with gaps between
ultimate branchlets flattened, covered with scale scales narrower than scales. Leaf gland absent
leaves. Leaves decussate, imbricate, dimorphic, with or very obscure; leaf apices of laterals obtuse 4
facials smaller than laterals, appressed, with apices 4a. Seed cones longer than wide when closed; seed
of laterals spreading above apices of facials; mar- wings nearly equal C.formosensis
gins entire; stomata inconspicuous, mostly on the 4b. Seed cones always globose; seed wings often
underside of the (leaf-covered) branchlets. Pollen unequal C. obtusa
cones terminal, solitary, very small, short cylindri-
cal; microsporophylls 816, decussate, bearing 23 Chamaecyparis formosensis Matsum., Bot. Mag.
abaxial, yellow or red pollen sacs. Seed cones termi- (Tokyo) 15: 137. 1901. Retinispora formosensis
nal, solitary, globose to ellipsoid-ovoid, small, with (Matsum.) A. V. Bobrov & Melikyan, Komarovia
peltate, parting scales. Bract-scale complexes (6)8 4: 74. 2006. Type: Taiwan: Nantou, Chia-i Pref.,
12(16), decussate, peltate, with a quadrangular, Yu-Shan, [Mt. Morrison], S. Honda s.n. (lectotype
rhombic or polygonal outline, abaxially depressed TI). Fig. 86, 87
with a small central umbo (bract tip), adaxially with
faint seed marks near base; the ultimate pair sterile. Etymology
Seeds moderately numerous, with 2 lateral, narrow
wings. Seedlings with 2 cotyledons. The species epithet indicates its origin as from
Formosa, an earlier name for Taiwan; perhaps
5 species. with a pun to Latin: formosus = handsome or well
formed.
Distribution
Vernacular names
North America: (disjunct) E North America; in
W North America in Oregon and California. Asia: Taiwan Cypress, Formosan Cypress; Beniki; hong
Japan, Taiwan. gui (Chinese)
Description Taxonomic notes
Trees to 5560 m tall; trunk monopodial, branch- This species may be more closely related to
ing low, occasionally forked, up to 67 m d.b.h. in Chamaecyparis pisifera of Japan than to its congener
a few very old trees. Bark soon flaky, on large trees on the island of Taiwan: C. obtusa var. formosana.
more than 20 cm thick towards base of trunk, fis- The two species in Taiwan, C. formosensis and C.
sured, fibrous, exfoliating in long strips, light red- obtusa, are sympatric but stay well separate taxo-
dish brown weathering grey. Branches spreading nomically. The much greater size of C. formosensis is
or curved down, higher order branches drooping, related to longevity and this is probably genetically
forming a dense pyramidal crown in young trees, determined; its seed cones are also larger and more
282 eventually becoming broadly conical or sympodial ellipsoid and the seed wings are more developed and
and domed in very old specimens. Foliage branches presumably more effective in dispersal as a result.
numerous, spreading and drooping; plagiotropic It is the largest conifer (measuring stem diameter)
branchlets alternating, smallest ones often unilat- growing in Asia and possibly the longest-lived as
eral on second highest order, gradually shortening well. A detailed study of its biology is very desirable.
forming tapering planate sprays, covered with green
leaves; ultimate lateral branchlets partly deciduous Distribution
after 46 years. Leaves decussate, imbricate, decur-
rent, scale-like, 13 0.51 mm on ultimate branch- Taiwan: central mountains.
lets, up to 10 mm long on leading shoots, dimorphic; TDWG codes: 38 TAI
facials smaller than laterals, rhombic to lanceolate,
carinate, obtuse-acuminate, appressed or with a Ecology
free apex, with an inconspicous abaxial gland; lat-
erals connate proximally, spreading above the apex In mixed coniferous forest of the cool temperate
of facials, conduplicate, lanceolate, recurved and coniferous forest belt between 1700 m and 2900 m
incurved at the mostly free apex, eglandular or a.s.l., with Chamaecyparis obtusa var. formosana
obscurely glandular; margins entire; leaves amphi- and C. formosensis as the dominant species, asso-
stomatic, stomata inconspicuous except for a few ciated with Calocedrus formosana, Cunninghamia
exposed lines on underside of facials; leaf colour konishii, and Taiwania cryptomerioides, in the upper
light grey-green or dull green. Pollen cones termi- part of the zone also Tsuga chinensis; in the under-
nal, solitary, ovoid-oblong, 23 11.5 mm, yellow- storey occur Photinia davidiana var. niitakaya
ish green turning brown; microsporophylls 812, mensis and the bamboo Yushania niitakayamensis,
decussate, peltate, suborbicular, with minutely den- but often little else. Chamaecyparis formosensis is
ticulate margins, with 3 abaxial yellow pollen sacs on exceptionally long-lived, estimated in excess of 3000
the lower margin. Seed cones terminal on branchlets years, and has a regeneration strategy similar to e.g.
with unmodified leaves, solitary, maturing within one Sequoiadendron, although the disturbance factors
year, caducous, subglobose to ellipsoid-ovoid, (6)10 are not well known. Soils are usually slightly acidic,
12 58 mm with opened scales, from purplish ripen- derived from igneous rock or shale. The climate is
ing to brown. Bract-scale complexes (8)1014(16), cool and wet, with abundant rainfall throughout the
decussate, parting and spreading at right angles from year and occasional snowfall in winter.
axis when mature, subpeltate to peltate, rhombic in
outline, 35.5 mm wide; abaxial surface depressed, Conservation
with a central recurved umbo (bract tip 0.50.7 mm);
margin undulating; base conical; adaxial face grooved The number of mature trees in natural forest stands
and striated, lustrous brown, lacking seed marks. has been greatly reduced by felling. The species
Seeds (1)2(3) at the basis of each scale (1520 occurs as scattered individuals in mixed coniferous
develop per cone), slightly flattened, oblong, 1.5 forest and has been selectively logged for its desirable
2.5 mm long, reddish brown, surrounded by 2 lateral, wood used in traditional oriental building. Growth
nearly equal thin wings 0.51 mm wide. is slow, especially in mature to over-mature phases
of its long life cycle. Several very large, senescent Description
individuals remain as natural monuments form-
ing a tourist attraction, but protection of more old Trees to 6070 m tall; trunk monopodial, large trees
growth forests containing this species is urgently can be buttressed at base, up to 44.5 m d.b.h. Bark
needed. The species is being grown in plantations. soon flaky, on large trees to 3540 cm thick towards
IUCN: EN (A2d) base of trunk, deeply fissured, fibrous, exfoliating
in long strips, dark red-brown weathering grey-
Uses brown. Branches slender, curved down; higher
order branches drooping or pendulous, forming
The wood (timber) of this species is highly prized a dense conical crown in young trees, eventually
for traditional oriental buildings such as temples becoming broadly columnar in large trees. Foliage 283
and shrines. It is uncommon in cultivation outside branches numerous, drooping or pendulous, plagio-
Taiwan, where this species is now being replanted in tropic branchlets alternating and smallest ones often
attempts to restore it from excessive logging in the unilateral, gradually shortening forming tapering
past. As an ornamental it is attractive, but slow grow- planate sprays, covered with green leaves; ultimate
ing and similar to Sawara cypress (C. pisifera) from lateral branchlets partly deciduous after 46 years.
Japan, which is much more common as a tree for the Leaves decussate, imbricate, decurrent, scale-like,
garden. To increase its use as an ornamental requires 23 11.5 mm on ultimate branchlets, up to 20 mm
marketing efforts and with so many cultivars already long on leading shoots, dimorphic; facials smaller
available from other species in the genus, this is not than laterals, rhombic to lanceolate, obtuse-acumi-
an easy task, and one which commercial growers nate, appressed, with a conspicous, oval, yellowish,
will be reluctant to undertake. Outside Taiwan, fine transparant, non-active abaxial gland; the laterals
specimen trees grow at Hilliers Arboretum in the connate proximally, spreading above apex of facials,
south of England, where ground frost is quite com- conduplicate, broadly falcate to lanceolate, recurved
mon; this species is apparently not as tender as is and incurved at the appressed apex, less conspicu-
commonly believed. ously glandular; margins entire; leaves amphisto-
matic, stomata inconspicuous, concentrated on the
partly hidden bases and margins of leaves; leaf colour
Chamaecyparis lawsoniana (A. Murray bis) Parl., light green or lustrous dark green; margins more or
Ann. Mus. Imp. Fis. Firenze 1: 181. 1864. Cupressus less glaucous, stomatal zones greenish white. Pollen
lawsoniana A. Murray bis, Edinburgh New cones very numerous, terminal, solitary, ovoid-
Philos. J., n.s., 1: 292. 1855; Retinispora lawsoniana oblong, 34 1.52 mm, yellowish green turning
(A. Murray bis) A. V. Bobrov & Melikyan, purplish black; microsporophylls 1216, decussate,
Komarovia 4: 74. 2006. Type: Illustration in peltate, with minutely denticulate margins, with 23
A. Murray, Edinburgh New Philos. J. n.s. 1, t. 10, abaxial large, red pollen sacs on the lower margin.
f. 14. 1855 (lectotype); USA: California, W. Murray Seed cones terminal on branchlets with unmodified
s.n. [3] (epitype E). Fig. 88, 89 leaves, solitary, maturing within one year, caducous,
subglobose, (6)711(14) mm with opened scales,
Etymology from glaucous-purplish (bracts) and yellow ripen-
ing to grey-brown. Bract-scale complexes 810(12),
This species was named after Charles Lawson (1794 decussate, ultimate pair often sterile and fused, part-
1873), an English nurseryman who grew several early ing and spreading at varying angles from axis when
introductions of conifers. mature, subpeltate to peltate, polygonal (45 angu-
lar) in outline, up to 67 mm wide, abaxial surface
Vernacular names slightly depressed, rugose, with a central small umbo
(bract tip 0.51 mm); base conical; adaxial surface
Port-Orford-Cedar, Port Orford White-cedar, grooved and striated, pinkish brown, with faint seed
Oregon Cedar, Lawson Cypress marks near the base. Seeds 24 at the basis of each
scale (1530 develop per cone), slightly flattened, was accidentally introduced in 1952, has led to a
ovoid, with acute apex, 2.53 2 mm long, yellow- steady decline over more than 150 years. Opening
ish, with 2 oblong, darker resin glands on each side, up areas formerly inaccessible with vehicles is part
surrounded by 2 lateral, nearly equal thin wings of the problem, as the pathogen gets transported by
11.5 mm wide. vehicles into new stands more quickly than would
otherwise have been possible. For this reason the
Distribution establishment of some 12 Research Natural Areas on
National Forest lands has been only partly effective
USA: SW Oregon, NW California, more extensively in halting the decline. Plantation forestry will largely
in Oregon (especially Coos Co., Curry Co.), from have to replace harvesting natural stands despite rel-
284 the Pacific coast to W of Mt. Shasta. atively slow early growth rates, in order to separate
TDWG codes: 73 ORE 76 CAL forestry from areas of infestation and to allow natu-
ral stands to recover. The major conservation effort
Ecology now undertaken concerns research to find and select
genotypes that are resistant to the pathogen causing
This species is usually growing in mixed conifer- root rot, to produce seed from these genotypes, to
ous forest of different types, with e.g. Abies con test the field performance of these genotypes on sites
color, A. grandis, Picea sitchensis, P. breweriana, with the pathogen present, and finally the reintro-
Pinus spp., Pseudotsuga menziesii, Sequoia semper duction of the resistant genotypes into the forests
virens, and Tsuga heterophylla, in various mixtures; where the species has been eliminated or reduced by
also in mixed conifer/ angiosperm forest with e.g. the disease.
Quercus chrysolepis and Lithocarpus densiflorus. IUCN: NT
Understorey vegetation is usually made up of the
ericaceous shrubs Rhododendron and Vaccinium, Uses
Rhamnus spp., Rubus spectabilis, Gaultheria shallon
and Quercus spp. and more locally Taxus brevifolia; The timber of this species is highly valued in East
a herbaceous flora is also commonly present. Except Asian countries, especially Japan, for construction
when growing with a dominance of Pinus, the forests of traditional buildings. There was a major inter-
are dense. Habitats less favourable to forest devel- national trade up to around 1990, especially with
opment, e.g. bogs and dry sand dunes, can still be Japan, where it fetched extremely high prices. There
occupied by Chamaecyparis lawsoniana, which will is probably no other conifer species which has given
then grow much slower and less tall in the commu- rise to more cultivar forms than C. lawsoniana, sev-
nities that prevail there. Best development is on wet eral hundred of which are listed in recent compila-
soil types with subsurface seepage, where the spe- tions (Leslie, 1992; Welch & Haddow, 1993). These
cies can become dominant; on drier sites the under lists were outdated upon publication as the number
lying rock is often ultramafic. Its altitudinal range is of new cultivar forms arising and sold by nurseries
from near sea level to ca. 1950 m a.s.l. The climate is seems to grow exponentially. Registration of these
strongly oceanic but summers tend to be warm and by the Conifer Registrar of the Royal Horticultural
dry (June-August); total precipitation ranges from Society in England, the acknowledged authority
10002250 mm annually, at higher altitudes a sub- in the world of conifer horticulture, has become
stantial part is snow. an almost impossible task. Many older cultivars
only exist, if at all, as one or a few surviving indi-
Conservation vidual shrubs or trees in gardens or parks and will
become extinct if not vegetatively propagated.
Over-exploitation of mature and old growth stands, Despite this, several cultivars are widely propagated
exacerbated with a more recent infestation by the and used, while the species itself is widely grown
oomycete pathogen Phytophthora lateralis, which from seed.
Chamaecyparis obtusa (Siebold & Zucc.) Endl., or dark brown. Bract-scale complexes 810, decus-
Syn. Conif.: 63. 1847. sate, parting and spreading at varying angles from
axis when mature; distal 2 connate and usually
Etymology sterile, peltate, polygonal (45-angular) in outline,
36 mm wide; abaxial surface depressed, rugose,
The species epithet refers to the obtuse (blunt) leaf with a central small umbo (bract tip 0.51 mm); base
apices. conical; adaxial surface grooved and striated, red-
brown, with light grey seed marks near base. Seeds
Vernacular names 2(3) at the basis of each scale (1520 develop per
cone), flattened, ovoid, with resin vescicles or longi-
Hinoki Cypress; hinoki (Japanese) tudinal grooves, 2.54 2 mm long, lustrous reddish 285
brown, surrounded by 2 lateral, unequal or nearly
Description equal thin wings 11.5 mm wide.
plate 11. Chamaecyparis obtusa var. obtusa. 1. Habit of tree. 2. Branch with foliage. 3. Branchlet with
leaves. 4. Juvenile leaves. 5. Adult leaves. 6. Branchlet with leaves and pollen cones. 7. Microsporophyll with
open pollen sacs and pollen. 8. Seed cones. 9. Seed cone scale. 10. Seeds.
and rain much of the year and very high precipita- pendulous; plagiotropic branchlets alternating and
tion exceeding 4000 mm per year. smallest ones often unilateral, gradually shortening
forming tapering planate sprays, covered with green
Conservation leaves; ultimate lateral branchlets partly deciduous
after 46 years. Leaves decussate, imbricate, decur-
Logging has caused a considerable decline in many rent, scale-like, coriaceous, 1.52 1 mm on ulti-
of the more accessible stands. After massive but mate branchlets, up to 15 4 mm on leading shoots,
unquantified destruction the total area of occu- dimorphic; facials smaller than laterals, rhombic to
pancy (AOO, in mixed stands with other conifers) obovate, sometimes obtusely keeled, obtuse-acumi-
50 years ago was ca. 50,000 ha for C. obtusa var. for nate, appressed, with a conspicous, circular, slightly
288 mosana and C. formosensis combined (Lee, 1962). It elevated, non-active abaxial gland; laterals connate
is almost certainly less than that now, as the decline proximally, spreading above apex of facials, condu-
of natural forest has continued from 66% to 52% of plicate, broadly falcate, recurved and incurved at
the land surface (Yang & Pan, 1996). Old growth for- appressed apex or convex on ultimate branchlets,
est is being replaced by secondary vegetation; where less conspicuously glandular; margins entire; leaves
active reforestation is undertaken the emphasis is amphistomatic, stomata concentrated on the under-
usually on faster growing conifers (Cryptomeria, side of laterals in depressed central region and on
Cunninghamia, Pinus). Important stands of partly hidden bases and margins of leaves; leaf colour
Chamaecyparis spp. are protected in the Yuanyang lustrous light or dark green, stomatal zones glaucous
Lake Reserve. white. Pollen cones numerous, terminal, solitary,
IUCN: VU (A2d) ovoid-oblong, 23 1 mm, yellowish green turn-
ing purplish brown; microsporophylls 812, decus-
Chamaecyparis pisifera (Siebold & Zucc.) Endl., sate, peltate, subcordate, with erose margins, with
Syn. Conif.: 64. 1847. Retinispora pisifera Siebold & 23 abaxial, yellow pollen sacs on the lower margin.
Zucc., Fl. Japon. 2 (5): 39, t. 122. 1844. Type: Japan: Seed cones terminal on branchlets with unmodified
Honshu, [loc. unknown], P. F. von Siebold s.n. leaves, solitary, maturing within one year, caducous,
(holotype not located, isotype K). subglobose or irregular, 57(8) mm with opened
scales, from yellowish green (bracts) and purplish
Etymology blue ripening to brown or blackish brown. Bract-
scale complexes (6)78(10), decussate, parting
The species epithet may refer to pear-shaped seeds. and spreading at varying angles from axis when
mature; distal pair usually connate, subpeltate to
Vernacular names peltate, quadrangular or irregular in outline, up
to 5 mm wide; abaxial surface centrally depressed,
Sawara Cypress; Sawara (Japanese) rugose, sometimes with a central small umbo (bract
tip 0.5 mm or invisible); base conical; adaxial sur-
Description face striated, light brown, with faint seed marks near
base. Seeds 12 at base of each scale (1015 develop
Trees to 4050 m tall; trunk monopodial, large trees per cone), slightly flattened, ovoid, 2 1.3 mm; apex
buttressed at base or not, up to 2 m d.b.h. Bark soon acute; brown, with 36 lighter resin vesicles on each
flaky, on large trees fissured, fibrous, exfoliating in side and a concave hilum at base; wings 2, lateral,
long strips, reddish brown weathering grey-brown. nearly equal thin, translucent, 1.52 mm wide.
Branches slender, ascending, spreading or curved
down; higher order branches spreading or droop- Distribution
ing, forming a dense conical crown in young trees,
eventually becoming broadly columnar in large Japan: Honshu, Kyushu.
trees. Foliage branches numerous, drooping or TDWG codes: 38 JAP-HN JAP-KY
Ecology Vernacular names
This species has a more scattered distribution in the Atlantic White-cedar, Atlantic White Cypress, White
forests than C. obtusa and is likely to be a palaeo- Cypress
relict. It usually occurs, where sympatric with the
more ubiquitous C. obtusa, on wetter sites e.g. near Description
mountain streams or in hollows with ground water
near the surface. In those wet places trees can spread Trees to 3035(40) m tall; trunk monopodial, up
by the layering of branches to form small groves of to 1.52 m d.b.h. Bark soon flaky, on large trees to
clonal individuals. It is a minor constituent in mixed 5 cm thick, deeply fissured and with connecting
conifer/angiosperm forests. Its altitudinal range is ridges, sometimes spirally twisted from torqued 289
from 280 m to 2600 m a.s.l. wood, fibrous, exfoliating in long strips, reddish
brown weathering grey-brown. Branches slender,
Conservation persistent, spreading or ascending; higher order
branches ascending, forming a conical crown in
Although being more scattered in its (original) dis- young trees, eventually becoming broader and more
tribution than C. obtusa, this species has not been open. Foliage branches numerous, slender, drooping
exploited anywhere nearly as intensively (its timber or pendulous; plagiotropic branchlets alternating,
value is limited) and, as a minor constituent, it sur- but smallest ones more irregularly disposed, only
vives where the natural forest containing it survives. slightly planated, forming tufts of foliage, covered
IUCN: LC with green leaves; ultimate lateral branchlets partly
deciduous after 45 years. Leaves decussate, imbri-
Uses cate, decurrent, scale-like, coriaceous, 1.52.5
11.5 mm on ultimate branchlets, up to 10 mm long
This species is limited as a source of timber, but on leading shoots, dimorphic, but facials on higher
widely cultivated as an ornamental, with many order branchlets only slightly shorter than the lat-
named cultivars. As of other members of the genus, erals, rhombic to ovate-oblong, sometimes keeled,
the wood is valued in Japan for traditional construc- obtuse or acuminate, appressed, with or without a
tion work, but its relative scarcity and the often poor raised, active abaxial gland in the centre; laterals
shape of layering trees limit its commercial use. connate proximally, spreading at or less often above
However, its ornamental merits are considered sub- the apex of facials, conduplicate, broadly falcate to
stantial due to the many cultivars with different hab- lanceolate, recurved and incurved at the appressed
its and foliage (among which are forms that retain apex, less conspicuously glandular; margins entire;
juvenile type leaves) selected in Japan and Europe. leaves amphistomatic, stomata inconspicuous, con-
Sarawa cypress is sensitive to drought and urban air centrated on the partly hidden bases and margins of
pollution, but is hardy in climates with not too long leaves; leaf colour yellowish green or greyish green.
and severe winters. Pollen cones numerous, terminal, solitary, ovoid,
1.53 12 mm, yellowish turning light brown or
dark brown to blackish brown; microsporophylls
Chamaecyparis thyoides (L.) Britton, Sterns & 812, decussate, peltate, subcordate, with minutely
Poggenb., Prelim. Cat. Anth. Pter. New York: 71. 1888. erose margins, with 2(3) abaxial large, yellow pol-
len sacs on the lower margin. Seed cones termi-
Etymology nal on branchlets with unmodified leaves, solitary,
maturing within one year, caducous, subglobose,
The classical Greek name for the Sandarac tree 47(8) mm with opened scales, from glaucous-
(Tetraclinis articulata) or its wood was thyon, hence purplish ripening to reddish brown or dark brown,
thyoides = similar but not equal to thyon. often glaucous. Bract-scale complexes (4)68(10),
usually 6 in decussate pairs, parting and spreading Burns & Honkala, 1990). However, the species is still
at varying angles from the axis when mature, pel- very widespread (though scattered) and common
tate, irregularly angular with curved margins, up in most swamp forests on the Atlantic Coastal Plain
to 35 mm wide; abaxial surface depressed, rugose, and Mexican Gulf coast in Florida and Alabama,
with a central curved, spiny umbo (bract tip ca. and is likely to have recovered in numbers if not yet
1 mm long); base narrowing; adaxial surface striated, in volume.
reddish brown, with faint seed marks near the base. IUCN: For ratings see under varieties.
Seeds 2 at base of each scale (812 develop per cone),
slightly flattened, ovoid, shallowly grooved, with Uses
concave light coloured hilum and acutish apex,
290 22.5 2 mm, lustrous dark brown, surrounded by The wood of this species is light, decay resistant, and
2 lateral, nearly equal wings 1 mm wide. is still widely used in the SE USA for many purposes
involving outdoor utilities. Trees are occasionally
Distribution cultivated and sold as ornamentals but there is no
substantial horticultural trade. A limited number
E and SE USA, from Maine south to N Florida, west of cultivars is known and most of the plants in cul-
to S Mississippi. tivation belong to one of these. Among these are
TDWG codes: 75 CNT MAI MAS MRY NWH NWJ some dwarf forms and one of these, Andelyensis
NWY RHO 78 ALA DEL FLA GEO MSI NCA SCA VRG has incorrectly been described as a botanical variety
by the German horticultural botanist Camillo Karl
Ecology Schneider. Forms originating in cultivation are not
botanical varieties (or species) and under the rules
Chamaecyparis thyoides usually grows in more of horticultural nomenclature are now to be given
or less pure stands in bogs and swamps and along non-latinized cultivar names.
streamside corridors of lowland rivers surrounded
by other tree species, which form the main forest 2 varieties are recognized:
types of the region where it occurs. Due to its great
latitudinal range it is associated with different species
from N to S. The majority of these are angiosperms, Chamaecyparis thyoides (L.) Britton, Sterns &
which also occupy the greater total area, associated Poggenb. var. thyoides. Cupressus thyoides L., Sp.
conifers are mainly Pinus spp. and Taxodium disti Pl. 2: 1003. Type: USA: [locality unknown], P. Kalm
chum. The soil types under stands of C. thyoides are LINN 1137.4 (lectotype LINN). Fig. 90
acid organic (muck) or sandy, with the water table
reaching the surface and prolonged seasonal periods Description
of inundation. It avoids salinity although it is known
to border tidal marshes in New Jersey. It is likely that Scale leaves on ultimate branchlets conspicuously
recurring fire would historically have been the dis- glandular, especially facials, these not keeled. Pollen
turbance agent preventing Acer rubrum from even- cones at maturity dark brown or blackish brown.
tually replacing C. thyoides in the succession. Apart
from the species composition of the vegetation due Distribution
to latitude, the ecology of the two varieties in this
species is similar. Eastern USA: coastal plain from Maine to Alabama.
TDWG codes: 75 CNT MAI MAS MRY NWH NWJ
Conservation NWY RHO 78 DEL FLA GEO NCA SCA VRG
This species has been heavily exploited for its timber Conservation
and total volume is considered to have been much
reduced during the 20th century (Little & Garrett in IUCN: LC
Chamaecyparis thyoides (L.) Britton, Sterns & Distribution
Poggenb. var. henryae (H. L. Li) Little, Madroo 18:
165. 1966. Chamaecyparis henryae H. L. Li, Morris SE USA: along the Gulf Coast from Florida to
Arbor. Bull. 13 (3): 43. 1962; Chamaecyparis thyoides Mississippi.
(L.) Britton, Sterns & Poggenb. subsp. henryae TDWG codes: 78 ALA FLA MSI
(H. L. Li) E. Murray, Kalmia 12: 19. 1982. Type: USA:
Florida, Escambia Co., Perdido River, at Barineau Conservation
Park, M. G. Henry 23 (holotype PH).
IUCN: LC
Description
291
Scale leaves on ultimate branchlets eglandular or
inconspicuously glandular, facials often distinctly
keeled abaxially. Pollen cones at maturity light
brown.
Cryptomeria D. Don, Ann. Nat. Hist. 1: 233. 1838. Type: Cryptomeria japonica
(Thunb. ex L. f.) D. Don [Cupressus japonica Thunb. ex L. f.] (Cupressaceae).
Greek: kryptos = covered, hidden; -meros = share 8 m) diam. Bark on large trees 23 cm thick, reddish
(parts or their number); referring to seeds hidden by brown, weathering grey, exfoliating in long, shred-
bracts. ding strips. Branches spreading to assurgent, form-
ing a conical crown in young trees and a rounded
Description crown in mature trees, self-pruning to leave a clear
bole in large trees. Foliage branches dense, shedding
292 See the species description. not individual leaves but ultimate lateral branches
which persist 48 years. Leaves helically arranged
Distribution in ranks of 5, decurrent, free for 1/23/4 of length,
spreading but incurved in various degrees, directed
As for the species. forward, linear-subulate, slightly flattened laterally,
distinctly keeled abaxially, stiff, green, 320(25)
12 mm; margins entire; apex acute; amphistomatic,
Cryptomeria japonica (Thunb. ex L. f.) D. Don, stomata in 4 greenish bands separated by green ribs,
Trans. Linn. Soc. London 18: 167. 1839. Cupressus usually on entire length of leaf. Pollen cones numer-
japonica Thunb. ex L. f., Suppl. Pl.: 421. 1781. ous, axillary and crowded towards ends of 2nd-year
Type: Japan: Honshu, [Habitat in Japonia], C. P. branchlets, 36 23 mm, elongating to 10 mm at
Thunberg UPS 22564 (lectotype UPS). Fig. 91 anthesis; microsporophylls 1530, the first 2 opposite
and green, then helically arranged, imbricate, pel-
Cryptomeria fortunei Hooibr., Wiener J. Gesammte tate, with 46 connate pollen sacs on the lower abax-
Pflanzenr. 1: 22. 1853, nom. nud. [fortunini]; ial margin. Seed cones terminal on down-curved
Cryptomeria japonica (Thunb. ex L. f.) D. Don var. branchlets with normal leaves, often aggregated or
fortunei (Hooibr.) Henry, in Elwes & Henry, Trees solitary, occasionally with proliferating vegetative
Gr. Brit. Ireland 1: 129. 1906 (nom. inval., Art. 34.1). short shoot at apex, globose to subglobose, squar-
Cryptomeria japonica (Thunb. ex L. f.) D. Don var. rose with spreading bract-scale complexes, soft
sinensis Miq., in Siebold & Zuccarini, Fl. Japon. 2: 52. woody, 1220(25) mm diam. Bract-scale complexes
1870; Cryptomeria japonica (Thunb. ex L. f.) D. Don helically arranged on a very short axis, 2540, con-
subsp. sinensis (Miq.) P. D. Sell, Watsonia 18 (1): 92. nate but parting at maturity, cuneate-rhombic, nar-
1990. rowly based; apex a curved, triangular, thin bract;
teeth adaxial to the bract below the upper margin of
Etymology the scale, (1)35(6) in number, acute to rostrate,
(2)2.55(6) mm long. Seeds (1)25 per bract-
The species epithet refers to Japan, the country of scale complex (some ovules may abort) depend-
origin of this species. ing on space available when intercalary scale tissue
develops, 45 3 mm, flattened, irregularly ovate
Vernacular names with 2 wings; wings unequal, 11.5 mm wide, form-
ing a strip around the seed.
Japanese cedar; sugi, omote-sugi, yaku-sugi,
(Japanese) Taxonomic notes
Named after Allan Cunningham (17911839), English Key to the species of Cunninghamia
traveller and botanist.
1a. Leaves 1230 23 mm, conspicuously amphi-
Description stomatic; adaxial stomata in continuous narrow
bands from base to apex of the leaf C. konishii
294 Evergreen, monoecious trees; trunk monopodial, 1b. Leaves 3060 35(6) mm, hypostomatic, or
readily resprouting. Resin ducts in leaves. Bark fis- at most with a few intermittent or incomplete
sured, fibrous, exfoliating in long strips, reddish lines of stomata adaxially C. lanceolata
brown. Branches in whorls, long, slender, higher
order branches spreading or drooping to pendulous,
plagiotropic (Massarts model); capacity to cop- Cunninghamia konishii Hayata, Gard. Chron., ser.
pice profound. Foliage branches slender, branch- 3, 43: 194. 1908. [J. Linn. Soc., Bot. 38: 299. 1908].
ing (nearly) opposite, lateral branchlets deciduous. Type: Taiwan: Chiayi Co., Luan-ta Shan, [Mt.
Leaves helically arranged, decurrent at distinctly nar- Rantaizan], N. Konishi s.n. (holotype TI). Fig. 92
rowed base, free part curved or twisted into a more
or less pectinate arrangement with primary stomatal Etymology
face on underside, coriaceous, flattened, linear-lan-
ceolate, gradually tapering to an acute apex; margins The species epithet commemorates N. Konishi, the
serrulate; amphistomatic or hypostomatic; abaxial collector of the type specimen.
stomata in 2 broad glaucous white bands separated
by a green elevated midrib; adaxial stomata in two Vernacular names
narrow bands (35 lines) along margins or absent.
Pollen cones subterminal on foliage branches, Taiwan shan mu (Chinese)
numerous in clusters subtended by a pseudo-whorl
of bract-like short leaves, cylindrical; microsporo- Description
phylls numerous, helically arranged, peltate, bearing
3(4) abaxial pollen sacs. Seed cones subterminal, Trees to 4050 m tall; trunk monopodial, often but-
solitary or paired, persistent (falling with foliage tressed in old trees, readily resprouting (coppice
branches); mature cones ovoid-globose. Bract-scale or pollard trees), to 3(4.5) m d.b.h. Bark on trunk
complexes numerous, helically arranged, imbricate, fissured, fibrous, exfoliating in long strips, reddish
appressed at base, spreading distally at maturity, brown weathering dull brown. Branches in whorls,
more or less triangular with a pedicellate base and long, slender, spreading to ascending near the top,
an apiculate to cuspidate apex, coriaceous; abaxial forming a pyramidal to finally irregular, rounded
surface smooth; adaxial surface with 23 seed marks crown; higher order branches spreading or drooping
at the distal end of the seed-bearing tissue. Seeds 23 to pendulous, 3rd and 4th order branches plagiotropic
per fertile scale, with 2 marginal, 1 mm wide wings. (but profuse reiteration after damage obscures this
Seedlings with 2 cotyledons. arrangement). Foliage branches numerous, slen-
der, branching (nearly) opposite, lateral branchlets
2 species. deciduous. Leaves decurrent at distinctly narrowed
base, free part curved or twisted into a more or less
Distribution pectinate arrangement with the primary stomatal
face on the undersides, flattened, narrowly lanceolate
China, Lao PDR, Taiwan, Viet Nam. or linear-lanceolate, straight or s-curved, gradually
tapering to an acute apex; margins serrulate, 1230 possible that we must conclude that it is a mon-
23 mm; amphistomatic, abaxial (lower) stomata tane form of a widespread, polymorphic species, or
in 2 broad glaucous white bands of 1020 irregular perhaps that we are dealing with a few cryptic
lines separated by a green elevated midrib; adaxial species.
stomata in two narrow bands (35 lines) along mar-
gins; adaxial surface smooth, glaucous green or Distribution
green. Pollen cones subterminal on foliage branches,
in clusters subtended by a pseudo-whorl of bract- Taiwan (N-central); China, Fujian; Lao PDR,
like short leaves, 1015 34 mm; microsporophylls Houaphan Province; Viet Nam (Bu Huong
50 or more, helically arranged on a slender axis, Mountain, Phu Hoat Mountain).
peltate; margins erose-denticulate; apex acute; bear- TDWG codes: 36 CHS-FJ 38 TAI 41 LAO VIE 295
ing 3 abaxial, large, oblong pollen sacs on the lower
margin. Seed cones subterminal, solitary or paired, Ecology
maturing within 1 year, persistent (falling with foli-
age branchlets); mature cones ovoid-globose, 1525 In mixed coniferous or conifer-broad-leaved forest
1520 mm, turning lustrous brown. Bract-scale com- in the cool temperate coniferous forest belt, with
plexes numerous, helically arranged on a thin axis, Chamaecyparis formosensis, C. obtusa var. formo
persistent, imbricate, appressed at base, spreading sana, Calocedrus formosana, Pinus taiwanensis,
distally at maturity, more or less triangular with a Pseudotsuga sinensis, Taiwania cryptomerioides,
pedicellate base, coriaceous; margins entire or den- Acer morrisonense, A. kawakamii, Schima superba,
ticulate near apiculate apex; abaxial surface smooth, Photinia davidiana, Rhododendron formosanum,
more or less keeled towards apex. Seeds usually 2 per Pasania sp., and the alpine bamboo Yushania niita
fertile scale, obovate, flat, 46 34 mm, brown with kayamensis. The altitudinal range is from (600?)1000
a light hilum near the base and 2 marginal wings m to 2200 m a.s.l. Soils are relatively deep and well-
1 mm wide leaving an emarginate seed apex. Teratism drained loams or loamy sand. The climate is cool,
in the form of shoot proliferation common. very moist, with cloud-cover resulting in fog and
rain much of the year and very high precipitation
Taxonomic notes exceeding 4000 mm per year.
Tassilicyparis A. V. Bobrov & Melikyan, Komarovia 4: Cotyledons 25, juvenile leaves only on seedlings, in
72. 2006. Type: Tassilicyparis dupreziana (A. Camus) whorls of 34, acicular-linear.
A. V. Bobrov & Melikyan [Cupressus dupreziana
A. Camus]. Platycyparis A. V. Bobrov & Melikyan, 15 species.
Komarovia 4: 73. 2006. Type: Platycyparis funebris
(Endl.) A. V. Bobrov & Melikyan [Cupressus fun Distribution
ebris Endl.]. Hesperocyparis Bartel & R. A. Price,
298 Phytologia 91 (1): 179. 2009. Type: Hesperocyparis SW North America, S to Honduras; North Africa
macrocarpa (Hartw. ex Gordon) Bartel [Cupressus and Mediterranean to Middle East; Himalaya to SW
macrocarpa Hartw. ex Gordon]. China.
Cupressus arizonica Greene var. revealiana Silba, Cupressus bakeri Jeps. subsp. matthewsii C. B. Wolf,
Phytologia 49 (4): 393. 1981. Aliso 1: 83. 1948.
Cuyamaca Cypress This species was named after M. S. Baker, who col-
lected the type specimen.
Description
Vernacular names
Bark on trunks smooth and exfoliating in thin flakes
and strips, exposing cherry-red bark, remaining Baker Cypress, Siskiyou Cypress, Modoc Cypress
smooth even on large trunks. Leaves variably glan-
dular, from eglandular on some branchlets to con- Description
spicuously and actively on (fewer) others, but mostly
inconspicuous. Boss on cone scales large or small. Trees 1015(25) m tall; trunk branching low, to
Seeds red-brown to dark brown, not or more or less 60100(140) cm d.b.h. Bark smooth, exfoliating
glaucous. with thin flakes exposing reddish, purplish red or
red-brown bark, giving a shaggy appearance, on
Distribution lower trunk to 2 cm thick and breaking into small
plates with upcurled edges, grey. Branches spread-
Mexico: Baja California Norte (Sierra Jurez), USA: ing, upper branches ascending, persistent, with foli-
S California (Cuyamaca Mts.). age towards end, giving a tufted appearance to the
TDWG codes: 76 CAL 79 MXN-BC often sparse, conical or pyramidal crown. Foliage
branches lax or rigid, ultimate branchlets opposite
Conservation or alternate, spreading at 3060 degrees, variable
in length (425 mm) but many ca. 10 mm, slender,
The two known populations of this variety are very 0.61.2 mm diam., quadrangular in cross-section,
restricted. Fires swept the upper King Creek drain- persistent. Leaves decussate, imbricate, on (sub)ulti-
age in 1950 and 1970 reducing the size of the popula- mate branchlets appressed, rhombic, 1.22 1 mm,
tion. A later fire in 2003 further destroyed most of keeled with a central depression; margins hyaline-
the remaining mature trees, but regeneration by seed denticulate or serrulate, obtuse or acute, on whip
shoots up to 10 34 mm, often with a spreading fields in which several of the populations occur. More
apex; stomata few near base abaxially, scattered from accessible populations are also threatened because
base to apex adaxially; glands conspicuous, central on National Forest fire suppression policy in the past has
rhombic leaves, producing resin droplets; leaf colour allowed the build-up of a dangerous fuel load, making
grey-green to dark green. Pollen cones terminal, sol- fires when they occur much more destructive.
itary, subglobose or ovoid, 23 2 mm, more or less IUCN: EN [B2ab (ii, iii, iv, v)]
4-sided; microsporophylls 812(14), decussate, pel-
tate, bearing abaxially 23(4) subglobose-angular Uses
pollen sacs. Seed cones terminal, solitary, grouped
but not densely clustered, maturing in two seasons No commercial uses are known of this species; like
304 to grey-brown or (silver-)grey subglobose cones of other cypresses trees may have been felled for fence
1018(22) mm diam. when slightly opened, persis- posts in the past. Although trees retain a conical
tent. Bract-scale complexes (4)68(10), decussate, crown which is attractive in gardens, it is little used
peltate, when growing with a prominent boss, when in horticulture, presumably because it is not strik-
full grown abaxially rugose, often verrucate, with a ingly distinct from other Californian species.
near-central, 23 mm long, triangular, or obscure,
umbo (bract tip), adaxially constricted to a narrow Cupressus cashmeriana Royle ex Carrire, Trait
base, grooved and ridged, dark brown, with lighter Gn. Conif., ed. 2, 1: 161. 1867. Type: United
seed scars. Seeds 4060(85) per cone, 34 3 mm, Kingdom: [cult. at the Royal Botanic Gardens, Kew
oval to triangular, more or less flattened, brown or (Himalayan House)], W. J. Bean & [?] Foster s.n.
red-brown, often glaucous, wings 2, forming thin, (neotype K).
narrow margins up to 0.5 mm wide.
Cupressus pendula Griff., Itin. Pl. Khasyah Mts.: 131.
Distribution 1848, non Thunb. (1783).
Cupressus tortulosa Griff., Not. Pl. Asiat. 4: 26. 1854
USA: N California (Siskiyou Co., Modoc Co., [et in Icon. Pl. Asiat. t. 372. 1854] [tortulosus or
Shasta Co., Plumas Co.), S Oregon (Jackson Co., torulosis, nom. ut. rej. (proposed 2010)].
Josephine Co.). Cupressus himalaica Silba, Phytologia 64: 80. 1987.
TDWG codes: 73 ORE 76 CAL
Etymology
Ecology
The species epithet means from Kashmir which was
In Pinus-Quercus woodland and, more commonly, an erroneous conjecture by the author of the spe-
above that zone in mixed conifer forest or wood- cies. [Under the Botanical Code, no name shall be
land with Pinus ponderosa, P. jeffreyi, Abies concolor, changed because it has the wrong meaning.]
Pseudotsuga menziesii, Calocedrus decurrens, Arcto
staphylos patula, and Seriphidium sp. (Artemisia); Vernacular names
predominantly on old lava flows in rocky or some-
times sandy soil. The altitudinal range is between Bhutan Cypress, Weeping Cypress
ca. 800 m and 2100 m a.s.l.
Description
Conservation
Trees to 8595 m tall (D. B. Gurung & S. Miehe,
This species occurs in 9 sites, some quite disjunct; unpubl. data); trunk often buttressed in large speci-
one of these comprises a population more than 3 km mens, to at least to 3.5 m diam. above the buttress.
across, the others are smaller or of unknown size, Bark becoming fibrous, reddish brown with pur-
and one population has become extinct in the 20th plish brown inner bark, exfoliating in long shaggy
century. Wildfires are the major hazard and difficult strips. Branches relatively long, spreading or ascend-
to fight due to the inaccessible terrain of the lava ing, often S-curved, ending drooping to pendulous,
forming a conical or pyramidal or in old trees based on young plants cultivated in the Jardin des
irregular and broad, dense crown. Foliage branches Plantes, Paris. Having received these plants from
(long) pendulous or more rarely drooping-pendu- another horticulturist without sufficient evidence
lous, very slender, lax, ultimate branchlets alternat- of provenance, his statements about origin were
ing, distichous, pendulous, gradually decreasing in speculative. No original material survives, but of the
length towards tip, together forming planate fron- several competing names originally available for the
dose sprays, ultimate branchlets semi-deciduous. Weeping Cypress of Bhutan (C. pendula Griff. 1848
Leaves decussate, imbricate, decurrent (long decur- unfortunately being a later homonym of C. pendula
rent on whip shoots with spreading apices) dimor- Thunb. 1783 and C. tortulosa Griff. now proposed for
phic (monomorphic on whip shoots), scale-like, official rejection for being obscure as well as ambigu-
with facials slightly smaller than laterals, on ulti- ous in its original spelling), C. cashmeriana Carrire 305
mate branchlets 1.43 0.51 mm, on whip shoots could be traced back to 19th century cultivated plants
up to 12 2 mm; laterals conduplicate, curved, their in Kew that may have the same origin as Carrires
free part spreading at or below the apex of facials, plant (Farjon, 1994). This material is also conspecific
with incurved or more rarely spreading acute with Griffiths collections of C. pendula Griff. and
apex; facials partly covered by laterals, with acute, a neotype was therefore selected from it. Cupressus
appressed or free apex; margins entire; glands con- corneyana Carrire remains incertae sedis and its
spicuous on facials, inconspicuous on laterals; sto- usage in Flora of Bhutan (Grierson & Long, 1983) is
mata few, scattered primarily on margins near leaf incorrect. There are apparently very glaucous, less
bases; leaf colour green or glaucous green, some- glaucous, and non-glaucous forms with long, pen-
times very glaucous. Pollen cones solitary and termi- dulous or shorter, more drooping foliage branches.
nal on ultimate branchlets, oblong, 46 22.5 mm; Horticulturists tend to make much of this, but we
microsporophylls 1016, decussate, peltate with must realise that in almost all cases we are dealing
erose-denticulate margins and more or less acute with consciously or unconsciously selected plants
apex, bearing 4 abaxial globose pollen sacs near the and their mostly clonal offspring. Trees growing
lower margin. Seed cones solitary or in groups near in the wild in Bhutan have greener, less pendulous
upper ends of pendulous branches, occasionally in foliage, and C. pendula Griff. and C. cashmeriana
persistent clusters, terminal on leafy branchlets with Carrire as originally seen and described by these
unaltered leaves, maturing in 2 growing seasons, authors are probably planted selections (cultivars).
(sub)globose to ovoid, (10)1221 1019 mm., I see no merit in recognizing the form in nature as a
green or glaucous green maturing to brown with distinct species C. himalaica Silba (e.g. in Grimshaw
parting scales. Bract-scale complexes 810 in decus- & Bayton, 2009: 299300).
sate pairs, of more or less equal size except for smaller
connate proximal pair, peltate, 45-angular in out- Distribution
line, centrally depressed with protruding 1 mm long
bract tip, rugose, abruptly narrowing towards the Indigenous in Bhutan, NE India: Arunachal Pradesh;
cone axis; adaxial surfaces striated, (red-)brown with planted widely in the region near Buddhist mon-
light grey seed marks near base. Seeds 1015 on each asteries and temples in E Nepal, Sikkim, Bhutan,
scale (fewer on the proximal pair), closely packed, Xizang [Tibet], and Arunachal Pradesh.
angular-ovoid, slightly flattened, ca. 4 2.5 mm, TDWG codes: 40 EHM-AP EHM-BH
reddish brown with whitish concave hilum at base,
wings 2 on opposite sides, unequal in size and shape, Ecology
11.5 mm wide to nearly absent.
A very large emergent in evergreen angiosperm for-
Taxonomic notes est dominated by Quercus, with lauraceous trees in
the understorey; also with Tsuga dumosa near the
The taxonomy and nomenclature of this species upper limit, and on rocky (limestone) cliffs in pure
have been confusing ever since Carrire (1855, 1867) stands. There are two possible strategies involved: late
described two new cypresses with slender, lax foliage successional stands depend on episodal disturbance
for regeneration, and extra-zonal avoidance of com- Vernacular names
petition on exposed cliffs. The altitudinal range is
from 1250m to 2670 m a.s.l. The climate in optimal Cheng cypress; min jiang bai mu (Chinese)
stands is strongly influenced by summer monsoon
rains, with ca. 8002000 mm annual precipitation. Description
China: S Gansu, N & W Sichuan. This variety is known from ca. 9 different locations
TDWG codes: 36 CHC-SC CHN-GS where natural populations are now mostly restricted
to more inaccessible sites such as canyons and exposed
Ecology cliffs. Overcutting is the main cause of its decline.
A collection from Hunan Province (L. B. Hu 1233, seen
In small, pure stands in some valleys, but more in E) is most likely from an introduced tree. Cypresses
commonly on rocky slopes or cliffs associated of this species are planted in villages within its natural
with Koelreuteria paniculata, Morus mongolica, range in Sichuan and possibly in Gansu as well.
Campylotropis delavayi, Bauhinia faberi, Cotoneaster IUCN: VU [B2ab (v)] 307
multiflorus, and C. gracilis; in non-acidic brown soils
over granites, quartzites and limestones. Based on Cupressus chengiana S. Y. Hu var. jiangensis
verified herbarium collections, the altitudinal range (N. Zhao) Silba, Phytologia 49: 394. 1981. Cupressus
is extensive, ranging from ca. 1200 m to 2750 m a.s.l. jiangensis N. Zhao, Acta Phytotax. Sin. 18 (2):
The climate is characterized by cold winters and 210. 1980, [jiangeensis]. Type: China: Sichuan,
cool to warm summers, with a distinct alternation Longmen Shan, Jiange Xian, L. S. Cai & T. Z. Min
of dry and rainy seasons; annual precipitation varies 101104 (holotype SCFI).
between 500750(1000 mm), with a 5070% mois-
ture deficit. There is no recorded difference in ecol- Description
ogy for the two varieties.
Seed cones ovoid-oblong, more equal in size; bract-
Uses scale complexes 1012.
Description Etymology
Seed cones globose to ovoid-globose, with (8) The epithet means: of the funeral (Latin funus = 311
1012(11) bract-scale complexes; seeds more or less death) and indicates its traditional use in China.
angular.
Vernacular names
Distribution
Funereal Cypress, Chinese Weeping Cypress; bai mu
Morocco (Oued-nFis valley). (Chinese)
TDWG codes: 20 MOR-MO
Description
Conservation
Trees to 3035 m tall; monopodial; trunk up to 2 m
The main population occurs in the Oued-nFiss val- d.b.h. Bark eventually becoming shallowly fissured,
ley between Asni and Ijoukak, where many old trees grey-brown, exfoliating in long strips. Branches rela-
still survive. Estimates indicate however that the tively long, spreading or ascending, ending droop-
total area of occupancy (AOO) suffered a reduction ing to pendulous, forming a conical or pyramidal
from ca. 5500 ha in 1950 to 1460 ha in 1986. More or in old trees irregular and broad crown. Foliage
recent surveys by Griffiths et al. in 1997 (unpubl. branches pendulous, slender, lax, ultimate branch-
exped. rep. Univ. of Reading, UK) failed to observe lets distichous, pendulous, forming planate frondose
natural regeneration even within an experimental sprays, slightly flattened (subtetragonous) in cross
enclosure designed to fence out the sheep and goats. section, ultimate branchlets semi-deciduous. Leaves
Exploitation for timber and firewood has certainly covering decussate, imbricate, decurrent, dimorphic
slowed down under current awareness programmes (monomorphic on whip shoots), scale-like, with
by the regional authorities, but protection is still facials slightly smaller than laterals, on ultimate
inadequate and further decline seems inevitable. branchlets 1.53.5 0.51 mm, on whip shoots up to
Afforestation to provide alternatives for local use of 10 2 mm; laterals conduplicate, straight or curved,
wood is needed, but great care must be taken with with appressed or more rarely free acute apex; facials
the risk of genetic introgression from closely related partly covered by laterals, with acute, appressed
species, e.g. C. sempervirens. apex; margins entire; glands on facials and laterals
IUCN: CR [A2acd; B2ab (iii)] (often more conspicuous on facials), linear; stomata
few, scattered primarily on margins near leaf bases;
leaf colour lustrous green. Pollen cones solitary
and terminal on ultimate branchlets, ovoid-oblong,
35 22.5 mm; microsporophylls 1014, decus-
sate, peltate, bearing 4 abaxial pollen sacs near the
lower margin. Seed cones solitary or in groups on
pendulous branches, terminal, maturing in 2 grow-
ing seasons, caducous, (sub)globose when closed,
(8)1015 mm diam., maturing to dark brown with Conservation
parting scales. Bract-scale complexes 68(12) in
decussate pairs, of more or less equal size except the The wide distribution of this species cited in the
smaller connate proximal pair, (510 mm wide), pel- Chinese literature (including Flora of China 4: 67.
tate, polygonal (45(6)-angular) in outline; abaxial 1999) is to a large extent based on planted or intro-
face more or less umbilicate, with a central curved duced trees outside the indisputably wild popula-
boss surrounding the protruding bract tip, finely tions. Occurrence in natural forests is rare due to
rugose, abruptly narrowing towards cone axis; adax- widespread deforestation and alteration of natural
ial surfaces angular, striated, chestnut-brown with vegetation in much of central China. The possibil-
whitish seed marks at base. Seeds 35(6) on each ity of secondary establishment from cultivated trees
312 scale, closely packed, angular-ovoid, slightly flat- outside its original range makes evaluation of threat
tened, 3 2.5 mm, dark brown with whitish hilum at to wild populations of this species extremely difficult.
base; wings 2 on opposite sides, unequal in size and The extent (and indeed existence) of natural popula-
shape, depending on room for growth 1 mm wide to tions of C. funebris is unknown at the current time.
nearly absent on one side. IUCN: DD
This species has often been classified in the genus This species is widely planted in China as an orna-
Chamaecyparis on the basis of its flattened foli- mental, especially common in Buddhist temple
age (dimorphic facial and lateral leaves on foliage grounds. Its wood is also considered valuable due to
sprays) and small cones. The ontogeny of its seed its durability and traditionally it was used for coffins
cones, which mature in two years, and the higher for this reason. In Europe it is uncommon in culti-
number of ovules/seeds, which arise in two succes- vation and most of the older extant trees are based
sive rows axillary to each bract, place it in Cupressus. on seed collection by Ernest Wilson made in Hubei
The dimorphic leaves are also found in other taxa in Province, China, in 1907. In horticulture it is often
the family adapted to higher athmospheric moisture, referred to as Chamaecyparis funebris due to the flat-
and are likely the result of convergent evolution. tened foliage and small cones.
In mixed mountain forest or (degraded) wood- Gowen Cypress, Mendocino Cypress, North Coast
land associated with Platycarya strobilacea, Vitex Cypress, Pygmy Cypress, Dwarf Cypress, Santa Cruz
negundo, Ligustrum sp., Viburnum sp., Pittosporum Cypress
sp., Myrsine africana, and Vitex negundo; in calcare-
ous soil or in sandy loam over sandstone; also widely Description
planted and probably invading into disturbed veg-
etation locally. The altitudinal range of this species is (Dwarf) trees to 10(50) m tall; monopodial; trunk
between 300 m and 2260 m a.s.l. to 1(2.4) m d.b.h. Bark eventually becoming
fibrous, greyish brown to grey, hard, fissured, with Ecology
anastomosing ridges, exfoliating in shreddy strips,
to 23 cm thick. Branches ascending or spread- In chaparral, pine barrens, and open pine wood-
ing, short or long, slender, often sparse, forming a land with Pinus attenuata, P. contorta, P. muricata,
conical or pyramidal crown, or in certain condi- P. ponderosa, P. radiata, Pseudotsuga menziesii,
tions dwarfed. Foliage branches spreading, slender, Arctostaphylos, Quercus, and Rhododendron, often
ultimate branchlets irregularly disposed, spread- in groves of up to 1000 trees or more; on sandstone
ing, short (35 mm) or longer (1020 mm) and outcrops, white or yellow sandy slopes, and leached,
rigid forming dense tufts, subterete in cross section, virtually sterile sandy hardpan, where it becomes
11.5 mm diam., torose, persistent. Leaves covering dwarfed. The altitudinal range is from near sea level
branchlets decussate, imbricate, decurrent on whip to 1200 m a.s.l. The climate is of the Mediterranean 313
shoots and then with recurved, free distal parts, on type with dry, hot summers, but in a narrow coastal
ultimate branchlets appressed, monomorphic, scale- strip cooled by frequent fog, and winter rain.
like, with facials equal in size to laterals, on ultimate
branchlets 11.5 11.3 mm and rhombic-gibbous, Uses
on whip shoots up to 5 mm long, with incurved and
appressed obtuse or more or less apiculate apex, Although introduced by C. T. Hartweg to England
sometimes slightly keeled; margins minutely dentic- in 1848, this species soon turned out to be tender
ulate; glands inconspicuous; stomata few, scattered in NW Europe and its cultivation outside collec-
primarily on margins near leaf bases; leaf colour yel- tions ceased. In southern Europe it is grown more
lowish green or dark green. Pollen cones numerous, widely in gardens and parks and a few cultivars are
solitary and terminal on ultimate branchlets, ovoid- known, some with doubtful affinity to this species
oblong, 34 1.52 mm; microsporophylls 1214, (see Vidakovi, 1991: 193).
decussate, peltate, bearing 35(6) abaxial pollen sacs
near lower margin. Seed cones in groups or clusters 2 varieties are recognized:
on lateral branches, terminal, maturing in 2 grow-
ing seasons, persistent, (sub)globose when closed,
1330 1225 mm, maturing to lustrous brown Cupressus goveniana Gordon var. goveniana.
and finally grey-brown or grey with parting scales. Callitropsis goveniana (Gordon) D. P. Little,
Bract-scale complexes 610, usually 8, in decus- Syst. Bot. 31 (3): 473. 2006, (nom. ut. rej., Art.
sate pairs, of more or less equal size, 815(18) mm 56); Hesperocyparis goveniana (Gordon) Bartel,
wide, peltate, polygonal (46-angular) in outline, Phytologia 91 (1): 181. 2009. Type: USA: California,
nearly flat or more often bossed, with protruding Monterey Co., Huckleberry Hill, [discovered
bract tip, abruptly narrowing towards cone axis; by Mr. Hartweg on the western declivity of the
abaxial surface rugose; adaxial surfaces striated, mountains of Monterey in Upper California],
brown or red-brown with whitish seed marks near C. T. Hartweg s.n. (holotype K).
base. Seeds 810 on each scale, closely packed, usu-
ally angular, slightly flattened, 35 24 mm, (black-
ish) brown or glaucous-brown with whitish hilum Cupressus goveniana Gordon var. pigmaea Lemmon,
at base; wings 2 on opposite sides, unequal in size Handb. W. Amer. Cone-bearers, ed. 3: 77. 1895;
and shape or discontinuous, depending on room for Cupressus pigmaea (Lemmon) Sarg., Bot. Gaz.
growth to 1 mm wide. (Crawfordsville) 31: 239. 1901; Cupressus goveniana
Gordon subsp. pigmaea (Lemmon) A. Camus,
Distribution [Les Cyprs] Encycl. Econ. Sylvicult. 2: 50. 1914,
[pygmaea]; Callitropsis pigmaea (Lemmon) D. P.
USA: California, Mendocino, Sonoma, Santa Cruz, Little, Syst. Bot. 31 (3): 474. 2006, (nom. ut. rej., Art.
San Mateo and Monterey Counties. 56); Hesperocyparis pigmaea (Lemmon) Bartel,
TDWG codes: 76 CAL Phytologia 91 (1): 182. 2009 (pygmaea).
Description Bonnie Doon is partly in an area with private real
estate development and fire prevention here will
Seed cones 1320 mm long; seeds (blackish) brown, allow pines like Pinus attenuata and P. ponderosa to
not or only slightly glaucous. crowd out the cypresses. The main population here
is within the Bonny Doon Ecological Reserve, sev-
Distribution eral others remain unprotected.
IUCN: CR [B2ab (ii, iii, v)]
USA: California, Mendocino, Sonoma and Monterey
Counties. Cupressus guadalupensis S. Watson, Proc. Amer.
TDWG codes: 76 CAL Acad. Arts 14: 300. 1879.
314
Conservation Etymology
Only known from a few scattered locations in a lim- The species epithet refers to Guadalupe Island, where
ited area, some of which are very close to continuing this species (var. guadalupensis) is indigenous.
urban development.
IUCN: EN [B2ab (ii, iii, v)] Vernacular names
Cupressus goveniana Gordon var. abramsiana Guadalupe Cypress, Tecate Cypress, Forbes Cypress;
(C. B. Wolf) Little, Phytologia 20: 435. 1970. ciprs (Spanish)
Cupressus abramsiana C. B. Wolf, Aliso 1: 215. 1948;
Callitropsis abramsiana (C. B. Wolf) D. P. Little, Description
Syst. Bot. 31 (3): 473. 2006, (nom. ut. rej., Art. 56);
Hesperocyparis abramsiana (C. B. Wolf) Bartel, Large shrubs or small trees 1015 m tall, rarely to
Phytologia 91 (1): 180. 2009. Type: USA: California, 20 m; trunk short, branching with several leaders
Santa Cruz Co., Bonnie Doon, ca. 1 km E of public low above ground, or sometimes monopodial with
School, C. B. Wolf RSA 6235 (holotype RSA). an unbranched trunk 23 m tall, to 1.21.5 m diam.
Bark remaining smooth and thin (to ca. 1 cm) even
Description on lower part of large trunks, exfoliating in small
flakes, or occasionally in long strips remaining partly
Seed cones (15)2030 mm long; seeds brown, often attached, exposing reddish or purplish brown to light
glaucous. chocolate-brown bark; outer bark weathering whit-
ish grey. Branches numerous, long, thick, spreading,
Distribution ascending or erect, forming in free standing trees
with short trunks a wide and rounded, but in more
USA: California, San Mateo and Santa Cruz crowded conditions a narrower or irregular crown.
Counties. Foliage branches lax or rigid, ultimate branch-
TDWG codes: 76 CAL lets 315 mm long, slender or thicker, 1.21.7 mm
diam., quadrangular in cross-section, becoming
Conservation thicker and subterete on older branchlets, covered
with leaves. Leaves decussate, imbricate, on ultimate
Restricted to four populations in the Santa Cruz branchlets appressed, rhombic, 11.5 1 mm, gib-
Mountains; two were located or found since the field bous or with a slight central depression; margins
studies by Wolf (in Wolf & Wagener, 1948). Some, hyaline-denticulate; apex obtuse, up to 15 34 mm
like the one near Eagle Rock, consist of only a few on some vigorous whip shoots; stomata on abaxial
score mature trees surrounded by highly flamma- side scattered in lower part, on the adaxial side from
ble chaparral, and/or are inaccessible to fire fight- base to apex; leaves eglandular (ultimate branch-
ers other than from the air. The population near lets) or with an inconspicuous, inactive gland,
covered with a thick cuticular wax layer, glaucous Uses
green to light green. Pollen cones numerous, termi-
nal, solitary, ovoid-oblong, terete or quadrangular No uses are known of this species and its varieties.
in cross-section, 47 23 mm; microsporophylls Only a few botanical collections in California (e.g.
1218(20), decussate, imbricate, peltate-cordate, Rancho Santa Ana Botanical Garden) have grown
abaxially bearing 34(5) angular-globose, yellow this species successfully. It should be taken into
pollen sacs. Seed cones terminal, not aggregated cultivation more widely especially for reasons of ex
in dense clusters but often grouped with up to 20 situ conservation of the variety native to Guadalupe
cones, growing in two seasons to globose or sub- Island.
globose cones, 1535 mm diam. (some subglobose
cones to 45 mm long), serotinous, coarsely rugose, 2 varieties are recognized: 315
sometimes with prominent bosses on the scales.
Bract-scale complexes (6)810, decussate, peltate,
45 sided or partially more or less rounded in out- Cupressus guadalupensis S. Watson var.
line, abaxially with a prominent and upcurved or guadalupensis. Callitropsis guadalupensis (S.
flat and inconspicuous boss near the centre, coarsely Watson) D. P. Little, Syst. Bot. 31 (3): 473. 2006,
rugose to verrucose, adaxially abruptly constricted, (nom. ut. rej., Art. 56); Hesperocyparis guadalupensis
dark brown with light seed scars towards base. Seeds (S. Watson) Bartel, Phytologia 91 (1): 181. 2009.
up to 100120 per cone, irregularly shaped, slightly Type: Mexico: Baja California Norte, Guadalupe
flattened, 47 23.5 mm, dark brown, the hilum Island, E. J. Palmer 92 (lectotype K).
conspicuously lighter; wings 2 narrow, irregular
strips on either side, 0.51 mm wide. Description
plate 12. Cupressus macnabiana. 1. Branch with foliage and seed cones. 2. Branchlet with leaves and
pollen cones. 3. Leaves with gland. 4. Microsporophylls with pollen sacs and pollen. 5. Seeds.
with a lighter 11.5 mm long hilum; wings 2, forming Vernacular names
very narrow strips around the seed.
Monterey Cypress, Ciprs Monterrey, Ciprs de
Distribution California (Spanish)
Cupressus torulosa D. Don var. torulosa. Cupressus Cupressus torulosa D. Don var. gigantea (W. C.
lusitanica Mill. subsp. torulosa (D. Don) Silba, Cheng & L. K. Fu) Farjon, Monogr. Cupressaceae
Acta Phytol. Yunnanica 28 (5): 470. 2006. Type: & Sciadopitys: 224. 2005. Cupressus gigantea W. C.
India: Himalaya, [Sooreh], W. S. Webb W 6046A Cheng & L. K. Fu, Acta Phytotax. Sin. 13 (4): 85.
(lectotype K-W). Fig. 99 1975. Type: China: Xizang [Tibet], [22 km from Jia-
mei-xi, Lang-bei Dong], Qinghai-Xizang Exped.
Cupressus austrotibetica Silba, Phytologia 65: 334. 3318 (holotype PE).
1988.
Cupressus karnaliensis Silba, J. Int. Conifer Preserv. Description
Soc. 1 (1): 19. 1994.
Cupressus karnaliensis Silba var. mustangensis Silba, Trees to 3040 m tall, up to 3.5 m d.b.h.. Ultimate
J. Int. Conifer Preserv. Soc. 1 (1): 22. 1994. branchlets short, usually thick, quadrangular to sub-
Cupressus tonkinensis Silba, J. Int. Conifer Preserv. terete in cross section, (1)1.52 mm diam., torose,
Soc. 1 (1): 23. 1994. persistent. Leaves on ultimate branchlets 11.8 0.7
Cupressus tongmaiensis Silba, J. Int. Conifer Preserv. 1.3 mm, rhombic-gibbous, glandular or eglandular;
Soc. 1 (1): 24. 1994. glands inconspicuous, in central depression, elliptic.
Distribution tinuous pressure for exploitation. Regeneration is
often poor due to grazing of livestock. Several groves
China: S Xizang [Tibet] (Zangbo River Valley, from are protected as sacred forest by Buddhist monks
ca. 93 to ca. 96 E); extreme NW Yunnan (vicinity of and some of the largest trees are on grounds used as
Dqn on the Langcang [Mekong] River). a cemetery, where any cutting is prohibited.
TDWG codes: 36 CHC-YN CHT IUCN: VU (A1d)
Conservation Uses
Due to the scarcity of timber in the region where Timber and firewood are locally taken. Large trees
326 scattered groves of var. gigantea occur, there is con- are often venerated in local traditions and religion.
Dacrycarpus (J. J. Bennett) de Laub., J. Arnold Arbor. 50: 315. 1969. Podocarpus sect.
Dacrydioideae J. J. Bennett in J. J. Bennett & R. Brown, Pl. Jav. Rar. 41. 1838. Type:
Dacrycarpus dacrydioides (A. Rich.) de Laub. [Podocarpus dacrydioides A. Rich.]
(Podocarpaceae).
Dioecious or rarely monoecious evergreen shrubs or Transitional forms between the two kinds of leaves
trees. Resin canals in leaves and seed cones. Bark hard, described in the key may exist in a single tree; it
occasionally with lenticels, becoming scaly. Leaves is therefore necessary to consider the extremes
trimorphic, with small scale leaves, acicular leaves of both forms where available in young trees and
and flattened, linear-falcate leaves, spirally inserted, mature trees alike. The two kinds are: scale-like,
decurrent at base, appressed to widely spreading; subulate or acicular (not wider than thick), and lin-
in some species distichous, flattened linear-falcate ear to S-curved, foliar (bilaterally flattened).
leaves on juvenile plants and on determinate vegeta-
tive lateral shoots of mature plants, but these absent 1a. All foliage branchlets with leaves of a similar
in other species. Small scale leaves on fertile shoots kind, shape acicular or subulate, sometimes
and on determinate vegetative lateral shoots in some slightly S-curved, 1.66(10) mm long, 0.41 mm
species; acicular leaves in the same positions in other wide 2
species, both leaf types not distichous, variously 1b. Leaves of different kind on different foliage
appressed to spreading. Stomata on both sides of the branchlets at least in young trees: scale-like,
leaves (leaves amphistomatic). Pollen cones single or subulate to acicular and linear to S-curved,
in pairs on axillary short shoots, at first nearly glob- bilaterally flattened and longer in young trees
ular but elongating to short cylindrical, to ca. 10 and with some species also both kinds in
3 mm; microsporophylls spirally attached to a slen- mature trees 4
der rachis, with triangular heads and two basal pollen 2a. Involucral leaves at base of seed cone 611 mm
sacs containing bisaccate pollen. Seed cones single long, incurved, enclosing the receptacle and
on axillary short shoots with scale leaves and often part of the seed(s) D. cinctus
surrounded by an involucrum of acicular leaves at 2b. Involucral leaves at base of seed cone 47 mm
ends of foliar branchlets, consisting of several spi- long, more or less straight, enclosing the recep-
rally arranged bracts; only a single subterminal one tacle only 3
becoming fertillized and forming an inverted seed; 3a. Leaves 1.63 mm long, acicular. Receptacles
the remainder becoming fused and partly swollen, 45 mm long, colouring dark purple; seeds
forming a verrucose receptacle ripening to red or 68 mm long D. compactus
purple and becoming succulent. Seeds completely 3b. Leaves 26 mm long, acicular to slightly
surrounded by the soft epimatium being partly fused S-curved. Receptacles 34 mm long, not chang-
with the fertile bract, forming a grooved or shallow ing colour or perhaps becoming yellowish;
crest on one side of apex. seeds 56(7) mm long D. expansus
4a. Pollen cones 2030 mm long; microsporophylls
9 species. subulate. Involucral leaves at base of seed cone
613 mm long, incurved, enclosing the recep-
tacle and part of the seed(s) D. cumingii
4b. Pollen cones 712 mm long; microsporophylls Etymology
apiculate. Involucral leaves at base of seed cone
18 mm long, spreading or incurved, at most The species epithet (Latin cinctus = enclosed or
only enclosing the receptacle 5 encircled) refers to the seed cones enclosed by invo-
5a. Pollen cones 710 mm long, 1 mm wide. lucral leaves.
Involucral leaves at base of seed cone 12 mm
long, leaving the 23 mm long receptacle free
Vernacular names
D. vieillardii
5b. Pollen cones 812 mm long, 23 mm wide. sareh (Sulawesi); djariha (and many other names,
Involucral leaves at base of seed cone 28 mm New Guinea)
328 long, at least in part enclosing the 37 mm long
receptacle 6
Description
6a. Involucral leaves at base of seed cone 58 mm
long, entirely enclosing the blue to purple Dioecious shrubs or trees to 30 m tall; trunk up to
receptacle D. kinabaluensis 1 m d.b.h., erect; branches contorted, spreading, crown
6b. Involucral leaves at base of seed cone 25 mm eventually more or less flattened. Bark exfoliating in
long, partly to nearly entirely enclosing the small plates or strips, brown weathering grey or black-
orange to red receptacle 7 ish. All foliage branches spreading to erect and with
7a. Involucral leaves at base of seed cone 23 mm leaves of similar kind, acicular and more or less bifa-
long, spreading, leaving most of the receptacle cially flattened. Leaves on all shoots spirally arranged,
free; seeds 3.54 mm long, smooth with a not or rarely somewhat distichous on terminal veg-
notched apex. Largest (foliar) leaves 37 mm etative shoots in young plants, all spreading slightly
long D. dacrydioides and equally around the shoot, acicular (thin and
7b. Involucral leaves at base of seed cone 35 mm hair-like on seedlings), curved inward, keeled abaxi-
long, incurved, enclosing part or nearly all of ally, 26(10) 0.40.8 mm, with curved, apiculate
the receptacle; seeds 47 mm long, grooved apex. Leaves of branchlets in crowns of mature trees
towards a curved apex. Largest (foliar) leaves shorter (24 mm) than on younger trees, spreading
(3)512(17) mm long 8 at ca. 45, decurrent, curved inward, apiculate, often
8a. Foliar, more or less S-curved, distichously glaucous. Stomata on all sides (leaves amphistomatic)
arranged leaves present in saplings and young but on abaxial side restricted to basal part of leaf, in 2
trees, 510 mm long, 0.81.1 mm wide. or more rows on adaxial face up to near apex. Pollen
Receptacles 34 mm long D. steupii cones terminal on short or long shoots, subtended
8b. Foliar, more or less S-curved, distichously by acicular leaves, nearly globose when immature, at
arranged leaves present both in young trees and maturity elongating to 810 mm long and 23 mm
in mature trees, (3)712(17) mm long, 12 mm wide. Microsporophylls with apiculate apex, ca. 1.5
wide. Receptacles 47 mm long D. imbricatus 0.6 mm, each bearing two protruding pollen sacs.
Seed cones terminal on short shoots with spreading,
Dacrycarpus cinctus (Pilg.) de Laub., J. Arnold 23 mm long, curved acicular leaves; involucral leaves
Arbor. 50: 332. 1969. Podocarpus cinctus Pilg., Bot. conspicuously longer, 611 mm, enclosing recepta-
Jahrb. Syst. 69: 253. 1938; Bracteocarpus cinctus cle and part of seed. Ripe receptacle 34 mm long,
(Pilg.) A. V. Bobrov & Melikyan, Bjull. Moskovsk. warty, red or purplish tinged. Ripe seeds 1(2) on a
Obsc. Isp. Prir., Otd. Biol. 103 (1): 58. 1998. receptacle, subglobose, 46 mm long, including the
Type: Papua New Guinea: Morobe, Bs River, smooth, light or dark red-brown, sometimes glau-
Mt. Sarawaket, M. S. Clemens 5261 (holotype B, cous epimatium, with a longitudinal grooved crest
destroyed; isotypes A, LAE, NY). Fig. 100 terminating in a curved protruding apex.
Distribution Etymology
330 New Guinea (highlands). This species was named in honour of the plant col-
TDWG codes: 43 NWG-IJ NWG-PN lector H. Cuming, who collected the specimens on
which the original description was based.
Ecology
Vernacular names
Dacrycarpus compactus is a highland species occur-
ring in subalpine shrubberies and on the fringes of sangu (Sumatera); igem (Phillipines: Davao,
alpine tussock grassland dominated by Deschampsia Mindanao) and other local names.
klossii. It is common in coniferous high montane
forest with Papuacedrus papuana, Podocarpus spp., Description
and a few angiosperms and then becomes more
abundant and often a dominant emergent tree in Dioecious trees to 38 m tall; trunk up to 1 m d.b.h.,
mossy low forest and shrubbery fringing peaty wet erect, monopodial; crown eventually spreading,
tussock grasslands. Here the deepest peat is grass- dome-shaped. Bark exfoliating in small plates or
land and the conifers Dacrycarpus and Papuacedrus strips, brown weathering grey. Foliage branches
occupy stony rises and scree slopes where the peat is with leaves of two kinds, more or less acicular and
thinner or absent and drainage better. Isolated trees flattened. Primary shoots of seedlings slender, of
may occur scattered in the grassland as they are rela- mature trees slender or in exposed crowns short and
tively resistant to grass fires once they reached some stiff, terminating in a bud-like cluster of incurved
size. There is a cool, misty climate at these heights acicular leaves. Leaves on all shoots spirally
that has almost no seasonal changes. The altitudinal arranged, subulate or acicular (thin and hair-like on
range of this species is (2800)30004300 m a.s.l. seedlings), curved inward at tip, spreading slightly,
keeled abaxially, with curved, apiculate apex. Leaves
Conservation on 26 cm long terminal and deciduous branchlets
of seedlings to young trees flattened, distichous,
IUCN: LC decurrent, linear or slightly s-curved, with parallel
smooth margins, 514 mm long (shortest at base and
Uses apex of branchlet), 0.81.3 mm wide, weakly keeled
on both surfaces; apex curved forward, apiculate.
This tree of moderate size produces useful timber of Leaves of branchlets in crowns of mature trees simi-
fine quality but its inaccessibility and relative scar- lar to more acicular, 24(6) 0.40.7 mm, less dis-
city of straight trees of good size prevent commercial tichously arranged to spreading on all sides, curved,
exploitation. The species is not in cultivation for for- apiculate. Stomata on both kinds of leaves and on
estry or horticulture. all sides (leaves amphistomatic) in many intermit-
tent rows on flat, larger leaves, in 2 or more rows on
each face of acicular leaves. Pollen cones terminal
on short shoots, subtended by small acicular leaves,
short when immature, at maturity elongating to Dacrycarpus dacrydioides (A. Rich.) de Laub.,
2030 mm long and 23 mm wide. Microsporophylls J. Arnold Arbor. 50: 337. 1969. Podocarpus
subulate like leaves, ca. 1.5 0.5 mm, each bearing dacrydioides A. Rich., in Lesson & Richard, Voy.
two protruding pollen sacs. Seed cones terminal on Astrolabe Bot. [Ess. Fl. Nouv. Zl.] 1: 358. 1832.
short shoots with spreading, 713 mm long, curved Type: New Zealand: North Island, J. S. C. Dumont
acicular leaves enclosing receptacle and seed. Ripe dUrville s.n., 1827 (holotype? G-DC). Fig. 101,
receptacle 34 mm long, warty, reddish. Ripe seeds 102, 103
1(2) on a receptacle, subglobose, 3.54.5 mm long,
including the smooth, light yellowish brown to Etymology
blackish brown, sometimes glaucous epimatium,
with a longitudinal grooved crest terminating in a The species epithet alludes to a similarity with 331
curved protruding apex. Dacrydium.
According to De Laubenfels var. patulus is restricted Dacrycarpus imbricatus (Blume) de Laub. var.
to Jawa, the Lesser Sunda Islands, and Sulawesi robustus de Laub., J. Arnold Arbor. 50: 323. 1969. 335
[Flora Malesiana ser. 1, 10 (3): 377, f. 25 (1988)] but Type: Papua New Guinea: Eastern Highlands, Mt.
collections at K and FHO that seem to belong to Wilhelm, L. J. Brass 30568 (holotype A).
this form with slender, inbricate primary leaves have
been gathered from Thailand to Vanuatu and Fiji, i.e. Podocarpus papuanus Ridl., Trans. Linn. Soc.
through the entire range of the species. The distinc- London, Bot., ser. 2, 9: 158. 1916; Bracteocarpus
tion between this form and branchlets with more papuanus (Ridl.) A. V. Bobrov & Melikyan, Bjull.
spreading primary leaves is often gradual and both Moskovsk. Obsc. Isp. Prir., Otd. Biol. 103 (1): 59. 1998.
may well occur in the same tree. Verification of spec- Dacrycarpus steupii de Laub., Kalikasan 7 (2): 127.
imens at KEP (Forest Research Institute of Malaysia) 1978, non de Laub. 1969.
and observations in the field in September-October
2008 confirmed this. Dacrycarpus imbricatus var. Description
patulus does not appear to be truly distinct and is
here treated as a synonym of Dacrycarpus imbricatus Leaves on primary shoots robust, 26 mm long,
var. imbricatus. 0.61 mm wide, often nearly all spreading around
the shoot, involucral leaves spreading.
Distribution
Distribution
As for the species.
Malesia: Borneo, Maluku [Moluccas], Philippines;
Conservation Papuasia: New Guinea.
TDWG codes: 42 BOR-BR BOR-KA BOR-SB BOR-SR
IUCN: LC MOL PHI 43 NWG-IJ NWG-PN
No common names are recorded for this species. Dacrycarpus kinabaluensis is a shrubby tree restricted
to the upper montane forest and subalpine dwarf
Description forest on Mt. Kinabalu (4101 m). It occurs on this
mountain from ca. 2600 a.s.l. m up to the tree line
Dioecious shrubs or small trees to 13 m tall, with at ca. 3500 m. Dacrycarpus imbricatus occurs to at
up to 30 cm stem diam.; trunk of trees short, often least 2400 m and possibly higher on this mountain,
gnarled; crown becoming dense. Bark exfoliating in but is a tree of taller forest. Dacrycarpus kinabalu
small plates or strips, brown weathering grey. Foliage ensis is growing predominantly on ultramafic rock
336 branches with leaves of two kinds only on seedlings but becomes one of the dominant shrubs above
and saplings, or on trees with adventitious shoots. 3000 m on granite. It can form dense, nearly pure
Leaves on primary shoots spirally arranged, more or stands but is commonly associated with other coni-
less imbricate, subulate or acicular, curved inward fers, e.g. Dacrydium gracile, D. gibbsiae, Phyllocladus
or more or less straight, keeled abaxially, 36 hypophyllus and Podocarpus brevifolius. Angiosperm
0.71 mm, with curved, apiculate apex. Leaves on trees are often scarce but heather-like tall shrubs and
24(5) cm long terminal (deciduous) branchlets of dwarfed trees are common with Rhododendron spp.
seedlings and saplings (or adventitious and shaded and Leptospermum spp. the most abundant. There is
shoots) flattened, distichous or nearly 4-ranked, often a thick moss layer on the forest floor, in which
decurrent, linear or slightly S-curved, with parallel orchids and pitcher plants (Nepenthes spp.) are com-
smooth margins, 510 mm long (shortest at base and mon, and epiphytes are abundant on the shrubs and
apex of branchlet), 0.81.3 mm wide, more or less low trees.
smooth; apex curved forward, apiculate. Leaves on
terminal branchlets in crowns of mature trees spi- Conservation
rally arranged or sometimes 4-ranked, never dis-
tichously arranged, shorter and narrower, curved, This species is only known from the higher parts
apiculate. Stomata on both kinds of leaves and on of Mt. Kinabalu and consequently occupies only
all sides (leaves amphistomatic) in 25 intermittent a small area probably less than 100 km in extent.
rows on flat, larger leaves, in 1 grooved row on each On the other hand, being in the centre of one of the
face of acicular leaves. Pollen cones terminal on short better protected National Parks in the whole of SE
shoots, subtended by small acicular leaves, nearly Asia provides it with a degree of protection, and no
globose when immature, at maturity elongating to decline is known at present. Increased tourism might
812 mm long and 23 mm wide. Microsporophylls pose indirect threats to this species, for instance
with triangular, apiculate apex, ca. 1.2 0.8 mm, by increasing the incidence of fires in episodes of
each bearing two protruding pollen sacs. Seed cones draught associated with El Nio climatic cycles.
terminal on short shoots with spreading, 58 mm IUCN: LC
long, incurved acicular leaves enclosing the recepta-
cle only. Ripe receptacle 45 mm long, warty, blue or Uses
purple. Ripe seeds 1(2) on a receptacle, subglobose,
57 56 mm long, including the smooth, light yel- This species occurs within a National Park at high
lowish brown to brown, sometimes glaucous epima- altitude and is not used economically. As far as
tium, with a longitudinal grooved crest terminating known it is not in cultivation except perhaps in a few
in a curved protruding apex. botanic gardens and/or private collections.
Distribution
341
figure 80. Cedrus libani var. libani in the Taurus Mts., Turkey
342
Dioecious or rarely monoecious evergreen shrubs The genus Dacrydium is one of the two earliest
or trees. Resin in bark and (1 canal) in leaves. Bark described genera in the family Podocarpaceae (the
hard, with numerous small lenticels, becoming other is Podocarpus) and it has in the past accomo-
fissured and scaly, exfoliating in plates. Primary dated numerous species, which are at present clas-
branches in pseudo-whorls, numerous or sparse, sified in other, in part segregate, genera, leaving a
spreading and assurgent, repeating the same pattern genus Dacrydium with a narrower, more precise
(Rauhs model), or with spreading and/or pendulous circumscription and at present containing 22 spe-
foliage branches; latter foliage branches not termi- cies. The segregate genera are, in order of publica-
nating in distinct buds. Leaves spirally arranged, tion date, Lepidothamnus (1860), Acmopyle (1903),
dimorphic, scale-like and subulate or acicular, usu- Falcatifolium (1969), Parasitaxus (1972), Halocarpus
ally curved inwards, gradually transitional from (1982), Lagarostrobos (1982) and, segregated again
juvenile to adult forms, keeled, triangular or quad- from Lagarostrobos, Manoao (1995). The circum-
rangular in cross-section, sometimes dorsiventrally scription of Dacrydium has remained open to various
flattened (i.e. wider than thick). Stomata on all sides interpretations until quite recently (Quinn, 1982), and
(leaves amphistomatic). Pollen cones solitary or in as a result conifer species that do not properly belong
groups, terminal or lateral on foliage shoots, axillary in the genus (even if including its segregates) or not
on short scaly peduncles or nearly sessile, becoming even in Podocarpaceae, have been ascribed to it. An
short cylindrical; microsporophylls peltate, triangu- informal classification into sections A, B and C by
lar to apiculate, with two basal pollen sacs contain- Florin (1931) divided the genus in its wider sense and
ing bisaccate pollen. Seed cones terminal on (short) only the species of his section B are still included in
foliage branchlets, small, consisting of several spi- Dacrydium. In subsequent accounts, when the genus
rally arranged, scale-like or leaf-like (acicular) bracts was circumscribed in a narrower sense, no attempts
following smaller scales; fertile bracts with a single were made to subdivide it (e.g. De Laubenfels, 1969,
more or less inverted ovule on the adaxial side; the 1988; Gaussen, 1974; Quinn, 1982). No phylogenetic
whole enlarged, swollen and red when ripe forming analysis of the genus has been published; at best a
a small receptacle with protruding bract tips in most few species were included in investigations into the
species. Seeds 12(3) per cone, becoming more or phylogeny of the family Podocarpaceae at the genus
less erect and usually standing obliquely from axis level. There is therefore no sound basis upon which
of subtending receptacle, protruding from a cup-like to base a classification of Dacrydium. Divisions of
epimatium covering its basal part only and ripening the genus based on a pre-selected few morphological
to dark lustrous brown or nearly black. characters would, as is explained under Podocarpus,
most likely be artificial. It is for this reason that this
22 species. relatively large (in conifer terms) genus must be
treated here in geographical groupings for the sake 4b. Adult leaves spreading 3045 from shoot; apex
of constructing identification keys that are manage- incurved but free and visible. Pollen cones
able and may work in the field. Few species are in 1.52 mm wide. Seed cones 23 mm long, with a
cultivation outside their native areas, so this should single, rarely 2 seeds D. balansae
not create undue problems. Two islands stand out as
logical entities for this purpose: New Caledonia and Key to the species of Dacrydium in Borneo
Borneo. The five species in New Caledonia are, as
the other conifers there, endemic. Most of the seven Juvenile leaf forms gradually merge into adult leaf
species of Borneo are also restricted to the island, forms and the extremes of both types should there-
but three species are also found elsewhere. These, fore be taken for comparison with the character
348 and the remainder (11 species) are dealt with in the states used in this key. Adult leaf forms are in most
third key. species shorter than juvenile leaf forms.
Key to the species of Dacrydium in New 1a. Adult leaves scale-like, appressed, 11.5 mm
Caledonia long D. elatum
1b. Adult leaves acicular, linear or linear-lanceo-
Juvenile leaf forms gradually merge into adult leaf late, spreading, (1.5)310(13) mm long 2
forms and the extremes of both types should there- 2a. Adult leaves and juvenile leaves similar, acicu-
fore be taken for comparison with the character lar, spreading wide to nearly 90, nearly straight,
states used in this key. Adult leaf forms are in most 510 mm long, 0.71 mm wide, flattened
species shorter than juvenile leaf forms. D. ericoides
2b. Adult leaves and juvenile leaves of different
1a. Shrubs or dwarfed trees up to 2.5 m (rarely to 4 length (adult leaves much shorter); adult leaves
m) tall. Juvenile and adult leaves similar in size spreading forward or in one species wide to 90
and shape or adult leaves only slightly shorter, 3
the shortest at least 5 longer than wide 2 3a. Adult leaves spreading wide to nearly 90,
1b. Small or large trees. Juvenile and adult leaves curved forward or nearly straight. Micro
very different in size and shape, the shortest less sporophylls of pollen cones with a rostrate apex
than 4 longer than wide 3 D. xanthandrum
2a. Adult leaves slightly shorter than juvenile 3b. Adult leaves spreading forward more or less
leaves, 510 mm long (juvenile leaves 1015 mm close to the shoot, usually curved. Micro
long), imbricate but with a free apex, narrowly sporophylls of pollen cones triangular or in one
lanceolate. Seeds 34 2 mm D. suprinii species lanceolate 4
2b. Adult leaves equally long as juvenile leaves, 4a. Adult leaves very slender and soft, 0.30.4 mm
(10)1315(17) mm long, spreading but curved wide, slightly curved or nearly straight
forward, acicular. Seeds 45 2.5 mm D. beccarii
D. guillauminii 4b. Adult leaves more or less rigid, (0.4)0.51.2 mm
3a. Adult leaves linear-lanceolate, 0.60.8 mm wide, curved at least towards the apex 5
wide; apex pungent. Pollen cones 47 mm long, 5a. Adult leaves robust, 0.91.2 mm wide, linear
1.2 mm wide D. lycopodioides or linear-lanceolate. Pollen cones 2025 4.5
3b. Adult leaves oblong, incurved apically, 11.8 mm 7 mm; microsporophylls lanceolate, 45 mm
wide; apex obtuse. Pollen cones 818 mm long, long D. gibbsiae
1.54 mm wide 4 5b. Adult leaves slender, 0.40.8 mm wide, acicular
4a. Adult leaves imbricate, strongly incurved api- or linear. Pollen cones 612 2 mm; microspo-
cally hiding the apex from view. Pollen cones rophylls triangular-apiculate, 11.5 mm long 6
34 mm wide. Seed cones 810 mm long, with 6a. Adult leaves linear, nearly as thick as wide, 0.4
13 seeds D. araucarioides 0.8 mm wide, keeled sharply on the abaxial
side; apex obtuse or short apiculate. Seed cones 6a. Shrubs or dwarf trees to 6 m tall; adult leaves
terminal on foliage shoots; epimatium covering crowded, spreading and curved forward,
the basal third of the seed D. pectinatum 1225(30) mm long, 0.61.2 mm wide
6b. Adult leaves acicular, wider than thick D. comosum
(transverse-triangular in cross-section), 0.40.5 6b. Trees to 35 m tall, sometimes shrubs at highest
(0.7) mm wide, keeled weakly on the abaxial altitudes; adult leaves spreading at ca. 90,
side; apex apiculate. Seed cones lateral on foli- straight or curved, (5)710(13) mm long, 0.5
age shoots; epimatium covering the basal half of 0.8 mm wide D. xanthandrum
the seed D. gracile 7a. Juvenile leaves 57 mm long, 1 mm wide, flat-
tened, 0.20.3 mm thick; adult leaves 24 mm
Key to the species of Dacrydium in regions long, 0.80.9 mm wide, flattened, 0.20.3 mm 349
outside Borneo and New Caledonia thick, linear-lanceolate D. spathoides
7b. Juvenile leaves 520(25) mm long, 0.20.6 mm
Juvenile leaf forms gradually merge into adult leaf wide, not flattened; adult leaves not flattened
forms and the extremes of both types should there- 8
fore be taken for comparison with the character 8a. Bracts subtending seed cones longer than the
states used in this key. Adult leaf forms are in most leaves immediately below these, 38 mm long,
species shorter than juvenile leaf forms. completely hiding the cones 9
8b. Bracts subtending seed cones shorter or equally
1a. Adult leaves 12(3) mm long, scale-like or long as the leaves immediately below these,
acicular, appressed or spreading forward and 13 mm long, exposing the cones and/or the
incurved, 0.20.5(0.7) mm wide 2 epimatium 12
1b. Adult leaves (1.5)310(30) mm long, acicular 9a. Juvenile leaves 814 mm long, strongly curved;
or linear-lanceolate, spreading forward or out- apex apiculate. Pollen cones 812 1.82 mm;
ward, if incurved mostly longer than 3 mm, microsporophylls triangular D. cornwallianum
0.31.2 mm wide 5 9b. Juvenile leaves 1020 mm long, straight or
2a. Adult leaves scale-like, appressed, on slender, slightly curved; apex pungent. Pollen cones
cord-like shoots 3 (7)1018 22.5 mm; microsporophylls apic-
2b. Adult leaves acicular, spreading forward and ulate or elongated 10
incurved, on slender, brush-like shoots 4 10a. Juvenile leaves very slender, 0.20.3 mm wide,
3a. Juvenile leaves 512 mm long, 0.40.6 mm not keeled abaxially. Microsporophylls of pol-
wide, keeled abaxially; bracts at base of seed len cones short, apiculate; seeds 3.54 mm long,
cones 3 mm long D. novo-guineense lustrous brown D. nidulum
3b. Juvenile leaves 1015 mm long, 0.3 mm wide, 10b. Juvenile leaves 0.40.6 mm wide, keeled abaxi-
keeled on 4 sides; bracts at base of seed cones ally. Microsporophylls of pollen cones elon-
1 1 mm, triangular D. elatum gated, with incurved apex; seeds 45 mm long,
4a. Adult leaves sharply keeled abaxially; apex brown or black 11
obtuse or sometimes acute D. nausoriense 11a. Adult leaves partly imbricate, straight with
4b. Adult leaves not keeled but transverse-triangu- incurved apex, 0.50.6 mm wide. Pollen cones
lar in cross-section; apex acute-pungent to acu- 79 2.5 mm. Bract scales subtending seed
minate D. leptophyllum cones 35 mm long D. medium
5a. Juvenile leaves 1533 mm long, 0.81 mm wide; 11b. Adult leaves free, spreading forward, incurved,
adult leaves (5)725(30) mm long, 0.51.2 mm 0.30.4 mm wide. Pollen cones 1016 22.5 mm.
wide 6 Bract scales subtending seed cones 68 mm
5b. Juvenile leaves 520 mm long, 0.20.6(1) mm long D. magnum
wide; adult leaves (1.5)210 mm long, 0.31 mm
wide 7
12a. Juvenile leaves 58 mm long, not much longer long, 0.71 mm wide and thick, keeled on four sides;
than adult leaves, keeled on 4 sides; adult leaves apex acute to apiculate. The (pen)ultimate branchlets
0.81 mm wide. Seed cones when fully devel- with adult leaves 48 mm thick; the leaves spirally
oped 68(10) mm long, swollen succulent and arranged, imbricate, spreading from base at <30
orange to red; seeds lustrous black and strongly incurved towards apex, oblong, 24
D. cupressinum (5) 11.5 mm, keeled from base abaxially, becom-
12b. Juvenile leaves 1020(25) mm long, much lon- ing convex towards the obtuse (invisible) apex, lus-
ger than adult leaves, keeled abaxially; adult trous green. Stomata on juvenile leaves on all sides
leaves 0.30.8 mm wide. Seed cones minute, ca. in lines between ribs (leaves amphistomatic), on
3 mm long; seeds lustrous brown 13 adult leaves largely confined to the adaxial side but
350 13a. Adult leaves (4)610 mm long, acicular, soft, a few near base abaxially. Pollen cones terminal or
0.30.4 mm wide. Seeds 12(3) per cone lateral on very short branchlets, cylindrical, 918
D. beccarii 34 mm; microsporophylls similar to adult leaves but
13b. Adult leaves (1.5)35(7) mm long, linear, smaller and shorter, nearly triangular with incurved
rigid, 0.40.8 mm wide; Only 1 seed per cone apex; pollen sacs not visible due to imbricate micro-
D. pectinatum sporophylls. Seed cones terminal, subtended by 3 mm
long leaves; bracts elongated to ca. 5 mm, less curved
than the leaves, covering the 810 mm long cone up
Dacrydium araucarioides Brongn. & Gris, Ann. to the seed, turning purplish red at maturity; epima-
Sci. Nat. Bot., sr. 5, 6: 244. 1866. Metadacrydium tium at base of seed. Seeds single or 23 per cone,
araucarioides (Brongn. & Gris) Baum.-Bod., partly covered by the bracts, ovoid with a slightly
Syst. Fl. Neu-Caledonien 5: 76. 1989. Type: New constricted apex, 44.5 mm long, brown.
Caledonia: Grande Terre, Province Sud, Canala,
[montagnes au-dessus de Canala], E. Vieillard
Distribution
1277 (lectotype P). Fig. 107, 108
New Caledonia: Grande Terre, Province Sud.
Etymology TDWG codes: 60 NWC
plate 13. Dacrydium beccarii. 1. Habit of tree. 2. Branch with foliage. 3. Leaf. 4. Branchlet with juvenile
leaves. 5. Branchlet with leaves and seed cones. 6. Seed cone with seed.
Dacrydium comosum Corner, Gard. Bull. Straits Ecology
Settlem. 10: 244, t. 10. 1939. Type: Malaysia: Malaya,
Negeri Pahang, Pine Tree Hill, E. J. H. Corner SFN Dacrydium comosum occurs on exposed mountain
33222 (holotype SING). Fig. 110, 111, 112 ridges as a local dominant in stunted mossy forest on
rocky acidic soil or shallow peat; in one area at alti-
Etymology tudes between 1170 and 1440 m a.s.l. (the altitudinal
range is incompletely known). Occasional individu-
The species epithet means bearing a tuft of leaves als may grow in forest below the ridge and attain tree
(Latin comosus = with much or long hair). size to reach the canopy.
No common names have been recorded for this Dacrydium comosum is known from two separate
species. areas and in total from less than five localities within
these areas. Corner (op. cit.) mentioned four trees
Description from the type locality, but the numbers at other
localities are not known. There is a risk of fire threat-
Small trees to 15 m tall, with a single stem to 50 cm ening these small populations coming from burning
d.b.h., on exposed ridges dwarfed and flat-topped. and clearing forest at lower elevations. Seed produc-
Branches spreading and ascending in candelabra tion seems to be low and seedlings are sparse.
fashion, making an umbrella-shaped crown with IUCN: EN [B1ab (iiv), B2ab (iiv)]
crowded, upturned foliage branches at ends of pri-
mary branches. Juvenile leaves and adult leaves very Uses
similar. Juvenile leaves spreading more perpen-
dicular to branch, acicular, curved forward, very No uses have been recorded of this species.
narrowly tapering, to 33 mm long, ca. 1 mm wide
at base, keeled abaxially, ending in a pungent apex. Dacrydium cornwallianum de Laub., Fl. Malesiana,
Adult leaves densely crowded and covering the shoot ser. 1, 10 (3): 366. 1988, [cornwalliana]. Type:
completely, spreading forward, acicular, straight or Indonesia: Papua, Danau Paniai [Wissel] Lakes,
slightly curving outward, 1225(30) mm long, 0.6 Arupa, C. Versteegh BW 3041 (holotype A,
1.2 mm wide at or just above a slightly widened base, isotype K).
flattened with a sharp abaxial keel, gradually taper-
ing to a pungent apex. Stomata on both sides (leaves Dacrydium nidulum de Laub. var. araucarioi
amphistomatic) in a few intermittent lines nearly to des de Laub., J. Arnold Arbor. 50: 293. 1969. Type:
apex. Pollen cones mostly lateral on foliage branches, Indonesia: Papua, Danau Paniai [Wissel] Lakes,
subtended by a few short leaves ca. 5 mm long, 810 Arupa, C. Versteegh BW 3041 (holotype L).
3 mm; microsporophylls terminating in a narrow
extension, 1.52 mm long and 0.5 mm wide at base, Etymology
with two basal pollen sacs. Seed cones few, terminal
on short foliage branchlets, ca. 3 mm long, nearly Although no explanation was given, the species epi-
hidden by leaves, with several lanceolate, coloured thet means of Cornwall.
bracts and 1(2) exposed, ovoid, light brown seeds
45 mm long covered at base by the epimatium. Vernacular names
The species epithet elatum means tall and refers to The morphological differences between Dacrydium
the large size some trees can attain. pierrei and D. elatum as stated by Hickel (op. cit.) and
Gaussen (1974) do not exist in the herbarium speci-
Vernacular names mens examined at K from Indochina. Among these
are 4 sheets of L. Pierre 1396 specifically mentioned
kayuru bukit, ru bukit (Malaya); ouk (Borneo); by Hickel for the alledged differences; one of these
sambinur (?) sheets (Kew barcode K000288637) is here desig-
nated as the lectotype of D. pierrei Hickel.
Description
Distribution
Trees of medium to large size, to 40 m tall; trunk
to 1 m d.b.h., much branched eventually developing S China: Guangxi; Indochina: Kampuchea [Cambo-
a broadly spreading dense crown, sometimes more dia], Lao PDR, Thailand, Viet Nam; Malesia: Borneo
or less flat-topped. Bark scaly; outer bark brown (Brunei, Sabah and Sarawak), Philippines, Peninsu-
to blackish; inner bark pink to red, exuding red lar Malaysia (including Penang Island), Sumatera
resin from slash. Foliage of two types, with juvenile (W coast).
leaves and with adult leaves. Juvenile leaves acicu- TDWG codes: 36 CHS-GX 41 CBD LAO THA VIE 42
lar, on seedlings and saplings very slender, 1015 BOR-BR BOR-SB BOR-SR PHI MLY-PM SUM
Ecology Vernacular names
Dacrydium elatum occurs in evergreen tropical rain- sempilor (in the Bintulu dialect, given with the name
forest from the lowlands up into the mountains at D. beccarii Parl. as the original identification of the
altitudes between 250 m and 2350 m a.s.l. It is often type collection)
found in wet places such as peat swamp forests inter-
mixed with patches of kerangas forest. In the latter Description
type of vegetation, which occurs on nearly white
sandy soil, conifers often dominate in the forest, with Trees 1018 m tall; trunk to 30 cm d.b.h., erect and
Agathis, Dacrydium and Podocarpus usually present. single-stemmed, branching to form a spreading
358 In more open situations in the border zone between crown. Bark scaly, brown; inner bark more or less
forest and swamp or savannah D. elatum is usually fibrous, reddish. Foliage branches mostly drooping
growing along streams. It appears to be restricted to or nearly pendulous, with more or less persistently
acidic soils derived from granite or sandstone, often juvenile type leaves becoming only slightly shorter
mixed with organic matter like peat. in the adult form on mature trees. Leaves linear,
straight, spreading at almost right angles to shoot
Conservation except on new growth, 510 mm long, 0.71 mm
wide, 0.20.3 mm thick, flattened but becoming
IUCN: LC slightly concave towards apex, sharply keeled on the
abaxial side, with a faint midrib adaxially, narrow-
Uses ing abrubtly at the distal end to an apiculate apex.
Stomata in two bands of 68 lines on adaxial side
Dacrydium elatum is probably the main source of from base to apex, only a few near base abaxially
sempilor wood in SE Asia, which includes spe- (leaves weakly amphistomatic). Pollen cones mostly
cies of Dacrycarpus, Dacrydium, Falcatifolium and lateral on foliage branches, subtended by reduced
Phyllocladus, between which the timber trade makes leaves, 710 mm long and 22.5 mm diam.; micro-
no distinction. It is a light and relatively hard wood sporophylls triangular, ending in a lanceolate apex
used in construction, window frames, doors, join- ca. 1 mm long, with two basal pollen sacs partly hid-
ery, furniture, flooring, interior finishing, veneer, den by the base of the microsporophyll. Seed cones
and plywood as well as pulp for the paper industry. mostly below end of foliage branchlets, terminal on
A volatile oil resembling cedar oil can be distilled very short dwarf shoots with reduced, 23 mm long,
from the wood, which is done during the pulping narrowly triangular leaves; cones 45 mm long; cone
process. This species is cultivated as an ornamental bracts 34 mm long, becoming reddish. Seeds one or
tree, mainly in countries within its natural range. two per cone; mature seeds not observed.
Taxonomic notes
Dacrydium ericoides de Laub., Fl. Malesiana,
ser. 1, 10 (3): 371. 1988. Type: Malaysia: Sarawak, De Laubenfels (1988) described this as a new species
Merurong Plateau, Bukit Skelap, E. F. Brunig, SFN in his treatment of the conifers in Flora Malesiana,
8722 (holotype L). distinguishing it from D. spathoides in having lon-
ger and straight leaves (as in juvenile plants) even on
mature trees. Only the fertile structures (both male
Etymology and female) have adult type (much reduced) leaves
at their base on very short dwarf shoots. The other
The species epithet refers to a supposed similarity species in Borneo with juvenile type acicular leaves
with Ericaceae (Heather family) or the genus Erica. has curved leaves which are not flat (not wider than
thick) terminating in an acute apex. Only a limited Description
number of herbarium collections are known, now
mostly at KEP in Malaysia. Small trees 212 m tall, usually monopodial but with
low, spreading and assurgent primary branches; foli-
Distribution age branches in dense tufts towards ends of primary
branches, forming a flat-topped or domed cande-
Borneo: Sarawak (Mt. Dulit, Bukit Lawi, Bukit labra crown. Juvenile leaves only on seedlings and
Skelap, Gunong Mulu, Gunong Murud). saplings, spreading widely or in young trees spread-
TDWG codes: 42 BOR-SR ing at 30 forward, covering shoot, 1218 mm long,
11.2 mm wide, straight or slightly curved, keeled
Ecology abaxially, pungent. Adult leaves robust, directed 359
forward, densely covering shoot, linear to linear-
This species is locally common in primary mossy lanceolate, (3)58(10) mm long, 0.91.2 mm wide,
forest on exposed mountain ridges between 1000 m sharply keeled abaxially; apex incurved, pungent to
and 2200 m a.s.l. apiculate. Stomata mostly on adaxial side in inter-
mittent lines from base to apex on both juvenile and
Conservation adult leaves, few near base on abaxial side. Pollen
cones terminal or on a short lateral branchlet, cylin-
This species is only known from five localities, where drical, 2025 4.57 mm; microsporophylls lanceo-
it is locally common. The type of forest it occurs in late, 45 mm long, 1.5 mm wide at base. Seed cones
and its habitat generally are not prime targets for terminal or often on short lateral branchlets with
logging, as the forest quality from that point of view smaller leaves; bract leaves longer, the upper one
is poor on exposed mountain ridges. subtending the seed nearly equally long as seed and
IUCN: LC spreading, leaving the seed exposed. Seeds solitary
or sometimes in pairs, obliquely positioned, ovoid,
Uses 45 mm long, covered for basal third by the epima-
tium, lustrous light brown.
No uses have been recorded of this rare species.
Distribution
Dacrydium gibbsiae Stapf, J. Linn. Soc., Bot. 42: 192,
t. 4. 1914. Type: Malaysia: Sabah, Ranau District, Borneo: Sabah (Mt. Kinabalu).
Mt. Kinabalu N.P., spur above Lobang, L. S. Gibbs TDWG codes: 42 BOR-SB
4162 (holotype K). Fig. 115
Ecology
Dacrydium beccarii Parl. var. kinabaluense Corner,
Gard. Bull. Straits Settlem. 10: 244, t. 9. 1939. This species occurs on Mt. Kinabalu from swampy
mossy forest at 15002000 m a.s.l. to high rocky
Etymology ridges and slopes up to 3600 m. It is mostly restricted
to the serpentine and its ultramafic erosion products
This species was named after Lilian Suzette Gibbs where competition from other trees is limited. Below
(18701925), who collected and studied the flora of the summit of Mt. Kinabalu it grows as a shrub on
Mt. Kinabalu. granite. The vegetation there is low and more or less
open, with other conifers, e.g. Dacrycarpus imbri
Vernacular names catus, Phyllocladus hypophyllus and Podocarpus
gibbsiae, and Myrtaceae, e.g. Leptospermum recur
No vernacular names have been recorded. vum, Tristaniopsis spp., Ericaceae and various small
oaks (Quercus) as codominants. At high altitudes and width. Juvenile leaves remotely set, spreading at
the common ericaceous shrub Rhododendron eri 5090 from shoot, acicular, very slender, 1220
coides has an uncanny resemblance to Dacrydium 0.30.4 mm, curved forward, triangular in cross sec-
gibbsiae until one sees the red tubular flowers. tion, pungent. Adult leaves more densely set, spread-
Codominant or common conifers that grow with ing at 3045 forward, closer to shoot on leading
D. gibbsiae at higher altitudes on the mountain are branches, acicular, slightly to strongly curved for-
Phyllocladus hypophyllus, Dacrycarpus kinabaluen ward, (3)46(8) 0.40.5(0.7) mm, triangular in
sis, and Podocarpus brevifolius. cross-section and slightly wider than thick, weakly
keeled abaxially, apiculate, dark green, flushing light
Conservation green. Stomata on all sides (leaves amphistomatic),
360 in 13 intermittent lines on each face to apex. Pollen
Although as far as is known this species is restricted cones terminal or lateral on branchlets with adult
to Mt. Kinabalu, where it is abundant in specific hab- leaves, 67 2 mm at maturity; microsporophylls
itats, no decline has been observed. Mount Kinabalu 0.7 mm wide at base, with extended narrow apex
is a National Park which receives many visitors who 0.61 mm long, with two basal pollen sacs, turn-
hike up the mountain, and fire prevention might ing brown. Seed cones mostly below ends of foli-
become an issue if tourism increases. Little is known age branches with adult leaves, terminal on dwarf
about the capacity of this species to regenerate after shoots, subtended by shorter involucral leaves, cov-
fire. ered with bracts 23 mm long and reaching to base
IUCN: LC of seed, turning reddish at maturity. Seeds usually
single, obliquely positioned, 33.5 mm long, covered
Uses for the basal half by the epimatium and turning lus-
trous dark brown to black.
No uses have been recorded of this species.
Distribution
Dacrydium gracile de Laub., Fl. Malesiana, ser. 1,
10 (3): 367. 1988, [gracilis]. Type: Malaysia: Sabah, Borneo: Sabah (Mt. Kinabalu and vicinity), one
Ranau District, Mt. Kinabalu N.P., Silau-Silau trail, locality in Sarawak.
D. J. de Laubenfels P 716 (holotype L). Fig. 116 TDWG codes: 42 BOR-SB BOR-SR
Etymology Ecology
The species epithet refers to the thin, slender (juve- Dacrydium gracile is a tree occurring scattered in
nile) leaves. lower montane rainforest, at altitudes between 950
m and 1800 m a.s.l. It is usually associated with the
Vernacular names conifers Agathis borneeensis, Podocarpus laubenfel
sii, Sundacarpus amarus, Falcatifolium falciforme,
No common names have been recorded for this Nageia wallichiana, and Dacrycarpus imbricatus on
species. soils poor in nutrients (kerangas forest); in Sarawak
it occurs in low canopy heath forest on sandstone.
Description
Conservation
Trees to 40 m tall; trunk to 100 cm d.b.h., usually
with a clear bole and with long, tortuous branches, This species is quite rare and occurs as individual
developing a broadly spreading and very open trees in primary forest. Most herbarium collections
crown, sometimes more or less flat-topped. Bark are from Mt. Kinabalu and vicinity (partly in Mt.
scaly; outer bark dark brown; inner bark exuding red Kinabalu National Park) and the disjunct locality in
resin from slash. Foliage of two types, with juvenile Sarawak (in a different habitat) could indicate that
leaves and adult leaves markedly differing in length it is more widespread than currently known. With
the present knowledge its area of occupancy (AOO) gated acicular apex, gradually shortening towards
is certainly less than 20 km2 but the total number of cone apex to 12 mm; pollen sacs 2, partly hidden
mature individuals is unknown. by the curved basal part of the microsporophyll.
IUCN: VU (D2) Seed cones terminal on long or short foliage shoots,
810 mm long, hidden by leaves. Seeds 15 per cone,
Uses proximally covered for a third to half by the epima-
tium and subtended by bracts; bracts long, leaf-like,
No uses have been recorded of this species; it can be much exceeding the seed; seeds ovoid-oblong, 45
assumed to be of value for its timber like other spe- 2.5 mm, brown, with a constricted apex.
cies in the genus that grow into tall forest trees. Its
protected status within Mt. Kinabalu National Park Distribution 361
makes exploitation at least of these trees unlikely;
outside this protected area it may be logged. New Caledonia (Grand Lac, Lac en Huit, Rivire des
Lacs).
Dacrydium guillauminii J. T. Buchholz, Bull. Mus. TDWG codes: 60 NWC
Hist. Nat. (Paris), sr. 2, 21: 282. 1949. Type: New
Caledonia: Grande Terre, Province Sud, Chtes de Ecology
la Madeleine, J. T. Buchholz 1728 (holotype ILL).
Fig. 117 Dacrydium guillauminii is a strictly riparian species
growing on the banks and shores of streams and
Etymology small lakes that are frequently flooded; its seeds ger-
minate in water-logged soil. The Plaine des Lacs is
This species is named after the French botanist a basin where small rivers and shallow lakes drain
Andr Guillaumin (18851974). ultramafic laterite soils and eroded serpentine hills,
and the water after heavy rains is often of a milky
Vernacular names colour (one locality is on the aptly named Pernod
Creek). This species does not occur above the high
No vernacular name has been recorded for this water line. The altitudinal range is 150275 m a.s.l.
species. It is commonly associated with Retrophyllum minus,
another podocarp restricted to this habitat. Just
Description meters away where the ground rises, the maquis
minier vegetation dominates; consequently the
Slender shrubs or dwarf trees to 2.5 m tall; stem thick- niche of this species is (literally) very narrow.
ened at base, often sparsely branched. Bark rough,
with horizontal stripes and large lenticels, breaking Conservation
into small scaly plates, brown weathering black-
ish grey; inner bark fibrous, red-brown. Branches The extreme rarity of this species and its very
spreading and ascending, with foliage branches restricted habitat alone put this species at risk of
upright and tufted near ends. Leaves permanently of extinction. The 23 populations are fragmented due
the juvenile type, densely covering branches, acicu- to the very local presence of its habitat; changes in
lar, curved forward or sometimes curved near base the water regime (e.g. by pollution or diversion of
only, (10)1315(17) 1 mm, transverse triangular streams caused by mining operations upstream)
in cross-section, pungent. Stomata in 25 intermit- could destroy these. Although one population is
tent lines on all three sides (higherst number of lines protected in the Chtes de la Madeleine Botanical
on adaxial side) from base to apex. Pollen cones both Reserve, the others are unprotected and on land
terminal and lateral; terminal cones 815 mm long under mining concessions (not actively mined at
and 4 mm wide when expanded; lateral cones at base present). The species is very sensitive to fire and
of a terminal cone and smaller. Microsporophylls numbers of individuals have declined, probably as
45 mm long near base of cone, with a greatly elon- a result of increased tourism; restoration of habitats
in the reserve is now being undertaken after tourism Taxonomic notes
became better regulated.
IUCN: CR [B1ab(iii)+2ab(iii);C2a(i)] This species is imperfectly known taxonomically as
no fertile material has ever been collected. Originally
Uses described as Podocarpus leptophyllus by Wasscher
(op. cit.) and transferred to Dacrycarpus by Gaussen,
There are no recorded uses of this species and (except it was finally settled in the genus Dacrydium by De
in a few botanical collections) it is not in cultivation. Laubenfels, where it undoubtedly belongs. The com-
bination under Dacrydium was first validly published
Dacrydium leptophyllum (Wasscher) de Laub. by Silba as cited above. Wasscher was uncertain of its
362 ex Silba, Phytologia Mem. 7: 27. 1984. Podocarpus placement for want of flowers and fruits and sug-
leptophyllus Wasscher, Blumea 4 (3): 414. 1941; gested it could belong in Dacrydium, but he unfor-
Dacrycarpus leptophyllus (Wasscher) Gaussen, tunately chose the wrong genus (which was then
Trav. Lab. Forest. Toulouse T. 2, 1 (2, 20): 150 (nom. treated in a much wider sense than at present). This
inval., Art. 33.4). 1974; Bracteocarpus leptophyllus remains a poorly known species of which few if any
(Wasscher) A. V. Bobrov & Melikyan, Bjull. specimens other than the type are identified.
Moskovsk. Obsc. Isp. Prir., Otd. Biol. 103 (1): 59.
1998. Type: Indonesia: Papua, Pegunungan Maoke, Distribution
Mt. Goliath, A. C. de Kock 39 (holotype BO).
New Guinea: Papua (Mt. Goliath).
Etymology TDWG codes: 43 NWG-IJ
Etymology Ecology
The species epithet refers to similarity of the foli- Dacrydium lycopodioides is an uncommon tree
age branches with Lycopodium, a genus of clubmoss in montane forest on serpentine, occurring in a
common in New Caledonia. few sheltered ravine forests or hollows which have
escaped the ravages of fire in the surrounding land- 363
Vernacular names scape. These forest pockets are naturally less prone to
burn than the drier woodland and maquis minier,
No vernacular name is known for this species. but are not absolutely safe from forest fires. This spe-
cies also occurs on moist mountain slopes and ridges
Description (e.g. Mt. Mou) where fires are infrequent. Here its
stature becomes more stunted to 510 m as it occurs
Trees to 25 m tall; trunk d.b.h. to 50 cm, erect, in towards the summit area at around 10001200 m
forest forming a clear bole; crown spreading, dense a.s.l. in association with the conifers Araucaria hum
and rounded in old trees. Bark scaly with small boldtensis, A. laubenfelsii (emergents), Falcatifolium
flakes, brown, weathering grey; inner bark more taxoides, and Prumnopitys ferruginoides in low forest
or less fibrous and light brown. Foliage in much with numerous angiosperms among which species
branched tufts, spreading to erect in mature trees. of the Myrtaceae are dominant.
Leaves on seedlings and juvenile trees acicular,
710 mm long, 0.20.3 mm wide and thick, spread- Conservation
ing at ca. 45 from shoot, slightly curved forward to
nearly straight, triangular in cross section, pungent. This species is known from several herbarium col-
Adult leaves shorter and gradually changing from lections made over a period of more than 150 years,
juvenile leaves, spirally arranged and spreading at some of which are without locality data. The locali-
3040, curved forward above base, linear-lanceo- ties recorded are Mt. Mou, Mt. Dzumac, Mt. Nakada,
late, (2)35(6) mm long, 0.60.8 mm wide just Mt. Humboldt, and a mountain ridge above the
above base and 0.30.4 mm thick, keeled sharply on Ouinn River; it was most frequently collected from
abaxial side, gradually tapering to a pungent apex. Mt. Mou. This may partly be due to relative ease of
Stomata on all sides (leaves amphistomatic), but access of the latter mountain and it may occur else-
extending to distal part of leaf only on adaxial side, where in the mountains of the SE part of Grande
in a few intermittent rows separated by the keel and Terre. It is obviously a relatively rare species, but it is
by a midrib on the adaxial side. Pollen cones termi- not at risk at present, thanks to the relatively undis-
nal or lateral below a terminal cone, 47 1.2 mm turbed mountain ravines in which it occurs.
when mature; microsporophylls narrowly triangu- IUCN: NT
lar, keeled abaxially, tapering to an acute apex, with
two basal pollen sacs. Seed cones terminal, often on Uses
dwarfed shoots, 23 mm long, subtended by short
(12 mm) leaves followed by longer (23 mm) flat- No commercial or other uses are recorded of this
tened cone bracts surrounding the basal epimatium, rare species.
becoming swollen and red at base. Seeds single, cov-
ered for only the basal third or less by the epima-
tium, 3.54 mm long, ovoid but slightly flattened,
lustrous brown.
Dacrydium magnum de Laub., J. Arnold Arbor. 50: from Malaya. Rather subtle differences in the adult
299. 1969. Type: Indonesia: Maluku, Obi, G. A. L. de foliage leaves lead however to a different appearance
Haan bb 23.806 (holotype L). of branchlets; the pollen cones of D. magnum are
longer and the two species occupy different habi-
Dacrydium beccarii Parl. var. rudens de Laub., J. tats, leading to greater tree size in D. magnum. The
Arnold Arbor. 50: 303. 1969. fact that the distributions of the two species do not
overlap is in the Malesian Archipelago of less signifi-
Etymology cance; disjunct distribution in this genus within spe-
cies and even varieties is rather commonplace, as it
The species epithet means large and was presum- is among other conifers.
364 ably given for its relatively large pollen cones.
Distribution
Vernacular names
Malesia: Maluku [Moluccas] (Obi Island);
No common names are recorded for this species. Papuasia: Louisiades Archipelago (Tagula Island),
DEntrecasteaux Islands (Normanby Island);
Description Solomon Islands (Choiseul).
TDWG codes: 42 MOL 43 NWG-PNSOL-SO
Trees to 30 m tall; trunk d.b.h. to 60 cm, monopodial,
crown compact, densely branched. Bark scaly, red- Ecology
dish brown; inner bark pinkish or reddish. Primary
branches spreading and ascending, terminating with Dacrydium magnum is a canopy tree in tropical rain-
dense tufts of upright foliage branches. Juvenile forest at altitudes between 60 m and 1200 m a.s.l.; on
leaves spreading widely, similar to adult leaves but ridges of mountains it can become the dominant for-
longer, to 20 mm, acicular, slightly curved, triangular est tree. On exposed sites it becomes more stunted,
in cross section, keeled abaxially, 0.40.5 mm wide; but not a low shrub.
apex pungent. Adult leaves slightly spreading but
strongly directed forward on shoot and more or less Conservation
imbricate, short acicular, (2)35(6) 0.30.4 mm,
mostly uniform on a shoot, curved with apex turn- The conservation status of this species is difficult to
ing back to the shoot (making it possible to strike assess due to its wide and scattered distribution. It is
a finger tip backwards over a foliage branchlet), tri- known from islands west and east of New Guinea but
angular in cross section, keeled abaxially, apiculate. not from New Guinea itself. It is undoubtedly a rare
Stomata on all sides (leaves amphistomatic) but more species which may well turn up in collections made
numerous on the adaxial side, in intermittent lines in New Guinea already or to be made in future. Due
to apex. Pollen cones terminal on foliage branchlets, to the possibly wider distribution than presently
1016 22.5 mm. Microsporophylls with elongated known, but at the same time the likelihood of exploi-
and strongly incurved apex and two basal pollen tation along with other species of Podocarpaceae
sacs. Seed cones terminal, ca. 3 mm long, subtended from which loggers will not distinguish it, it is here
by leaves similar in length and shape to adult leaves; listed as Near Threatened.
the cone bracts longer, 68 mm, spreading or curved IUCN: NT
inward partly covering the seed. Seeds solitary or in
pairs, obliquely positioned, covered for basal third Uses
by the epimatium, 45 mm long, ovoid, brown.
No uses have been recorded for this species.
Taxonomic notes
Trees of medium size, to 25 m tall; trunk to 50 cm This species exists in small stands in remnants of
d.b.h., much branched eventually developing a rainforest and is exploited for its timber. Much of
broadly spreading domed crown. Bark scaly with this forest is privately owned or on tribal land or
large flakes; outer bark brown weathering grey; inner both and there are no forest reserves where this spe-
bark pink to reddish. Foliage of two types, with juve- cies is being protected. It is mostly known from the
nile leaves and with adult leaves. Juvenile leaves acic- Nausori Highlands but other localities have been
ular, on seedlings and saplings very slender, 1015 found, one of them on Vanua Levu. Much of the for-
0.3 mm, shorter and slightly thicker on young trees, mer rainforest which was the habitat of this species
366 straight or slightly curved forward, spreading at has been converted to agriculture or pasture, espe-
more than 45 from shoot, keeled on four sides, cially in more densely settled areas. Regeneration is
pungent. Transitional leaves present on saplings, slow and does not appear to be adequate; this podo-
35 mm long, spreading at ca. 45. Adult leaves on carp probably needs periodic disturbance by fire to
numerous slender, much branching shoots, decur- spread from seed, but too frequent fires destroy that
rent at base, spreading out and forward at ca. 30 opportunity.
from shoot, short acicular, 11.5 mm long and 0.3 IUCN: EN [B2ab (iii)]
0.4 mm wide (on leading branches slightly larger),
keeled abaxially, obtuse or sometimes acute with Uses
incurved apex. Stomata on all sides (leaves amphi-
stomatic), on juvenile leaves in narrow grooves, on Considered a highly valuable timber tree in Fiji
adult leaves in a few scattered lines, absent on the (graded Class I), this species is used in construction
distal part of the abaxial side, on the adaxial side to and all kinds of carpentry work.
apex. Pollen cones terminal or lateral on branchlets
with adult leaves, 46 11.2 mm at maturity, micro- Dacrydium nidulum de Laub., J. Arnold Arbor. 50:
sporophylls triangular, 1 0.7 mm, acute-apiculate, 292. 1969. Type: Indonesia: Papua, Jazirah Doberai,
with two basal pollen sacs. Seed cones terminal on Lake Aja-Maroe, W side of lake [Segior], W. Vink
long or very short branchlets with adult leaves, ca. & M. Vink BW 15271 (holotype L).
3 mm long, with distal bracts 2 mm long, slightly
swelling and turning red at maturity. Seeds solitary, Etymology
free from leaves, obliquely positioned, covered for
basal third by the epimatium, ovoid, 3.54 mm long, The species epithet is derived from Latin nidus (=
turning lustrous brown. nest) and refers to the nesting of the growing seed
cone within its elongated leaf-like bracts.
Distribution
Vernacular names
Fiji (Vanua Levu, Viti Levu).
TDWG codes: 60 FIJ yaka (Fijian)
Ecology Description
Dacrydium nausoriense is a canopy tree in more or Trees to 30 m tall, monopodial and erect; trunk d.b.h.
less open montane rainforest (annual rainfall ca. to 70 cm or rarely more, usually with a clear bole;
3000 mm) and fairly common on the plateaux and crown spreading, dense and rounded in old trees.
summits of the Nausori Highlands. It is associated Bark irregularly fissured, scaly, dark brown, weather-
with Dacrycarpus imbricatus and Podocarpus neri ing grey; inner bark slightly fibrous and red-brown.
ifolius var. degeneri and with numerous angiosperm Foliage in much branched tufts, drooping to spread-
trees and shrubs. ing in mature trees. Leaves on seedlings and juvenile
trees long acicular, 1020 mm long, 0.20.3 mm wide Conservation
and thick, spreading widely, slightly curved to nearly
straight, triangular in cross section, pungent. Adult IUCN: LC
leaves much shorter but variable in length between
trees and within trees, spirally arranged and decur- Uses
rent at base, short acicular, (1.5)35(7) mm long,
0.30.7 mm wide and 0.30.4 mm thick, keeled In Fiji, where this species (and D. nausoriense, not
sharply on abaxial side, spreading at 3045, slightly distinguished by loggers) is considered a very valu-
curved or nearly straight but with an incurved, able timber tree, it is used mainly for construction
obtuse to short apiculate apex. Intermediate forms and carpentry such as furniture making.
of leaves occur on younger trees. Stomata on all sides 367
(leaves amphistomatic), but extending to apex only Dacrydium novoguineense Gibbs, Contr.
on the adaxial side, in several intermittent rows. Phytogeogr. & Fl. Arfak Mts.: 78. 1917. Type:
Pollen cones terminal or lateral, ca. 4 2 mm when Indonesia: Papua, Arfak Mts., Koebre Ridge,
immature, elongating to 18 mm; microsporophylls L. S. Gibbs 5648 (lectotype BM).
triangular, 1.2 0.8 mm, keeled abaxially, apiculate,
with two basal pollen sacs. Seed cones terminal, 23 Etymology
mm long, subtended by short (1.52 mm) leaves, fol-
lowed by longer (35 mm) cone bracts surrounding The species epithet refers to its occurrence in New
the epimatium, becoming swollen and red at base. Guinea.
Seeds solitary, covered for only the basal third or less
by the epimatium, 3.54 mm long, lustrous brown. Vernacular names
Ecology Description
Dacrydium novo-guineense is very common in New Small to medium size trees to 20 m, rarely to 40 m
Guinea, where it forms a canopy tree in open forest tall; trunk d.b.h. to 1.5 m or rarely more; crown
especially on mountain ridges. In forests at middle spreading, dense and rounded in old trees. Bark
altitudes (13002200 m) it is an emergent above the brown, weathering grey, peeling profusely in broad,
general canopy, but higher up on ridges it becomes a loose brown strips. Foliage in much branched tufts,
small tree among other scrub and tree ferns. It often drooping to more or less erect on mature trees.
regenerates abundantly after disturbance by fire, Leaves on seedlings and juvenile trees long acicular,
when it can become temporarily dominant. This 1020 mm long, 0.20.4 mm wide and thick, slightly
species is indifferent to soil types and occurs on clay, curved, abaxially keeled, pungent. Adult leaves much
sand, quartzite and other rock debris as well as on shorter but variable in length between trees and
peat. The altitude range of this species is very large, within trees, linear, (1.5)35(7) mm long, 0.40.8
from 50 m to 3000 m according to data from her- mm wide and thick, keeled sharply on abaxial side,
barium labels. spirally arranged and decurrent at base where they
are imbricate, spreading at 3045, slightly curved or
Conservation nearly straight but with an incurved, obtuse to short
apiculate apex. Intermediate forms of leaves present
IUCN: LC on younger trees. Stomata on all sides (leaves amphi-
stomatic), but extending to apex only on adaxial side,
Uses in several intermittent rows. Pollen cones terminal,
ca. 4 2 mm when immature, elongating to 12 mm;
The timber of this species is used for building and microsporophylls narrowly triangular, 1.5 0.5 mm,
the bark for insulation of the walls of traditional keeled abaxially, apiculate, with two basal pollen
houses in the highlands of New Guinea. sacs. Seed cones terminal, subtended by short leaves
followed by slightly longer (23 mm) cone bracts,
becoming swollen and red. Seeds single, covered for
only the basal third or less by the epimatium, ovoid,
45 3 mm, chestnut to lustrous dark brown.
Taxonomic notes Dacrydium spathoides de Laub., J. Arnold Arbor.
50: 299. 1969. Type: Indonesia: Papua, Pegunungan
This species resembles the lowland form of D. nidu Maoke, Idenburg River, 18 km SW of Bernhard
lum, but differs from it by its more robust leaves Camp, L. J. Brass 12659 (holotype A).
and by the fully exposed mature seed. The length of
leaves in mature trees is influenced by environmen- Etymology
tal conditions, with shorter leaves associated with
more adverse environments such as peat bogs. The species epithet refers to the shape of the adult
leaf resembling a spathe (as in palms) with its con-
Distribution cave adaxial side.
369
China: Hainan; Malesia: Borneo (including Karimata Vernacular names
and Serutu Islands), Philippines, Sumatera: Pulau
Bangka and Pulau Belitung. No common names have been recorded for this
TDWG codes: 36 CHH 42 BOR-BR BOR-KA BOR-SB species.
BOR-SR PHI SUM
Description
Ecology
Trees 2535 m tall; trunk monopodial, erect, to 50
Dacrydium pectinatum is a species of lowland to cm d.b.h. Bark scaly, brown weathering grey; inner
montane rainforests, where it occurs as scattered bark reddish and exuding red sap. Crown spread-
individual trees at altitudes from near sea level to ing above a clear bole in canopy forest. Juvenile and
1500 m a.s.l.; above 600800 m trees become scarce. adult leaves similar, mainly differing in size and with
More dense stands or even pure stands of this spe- a gradual transition. Juvenile leaves flattened acicu-
cies are restricted to nutrient-poor soils like shallow, lar, spreading at ca. 60, 57 mm long, 1 mm wide
leached sands or to swamps with peat formation and 0.20.3 mm thick, straight or slightly curved
above the water table. The forest on nutrient-poor forward at the pungent apex, keeled on abaxial
sandy soils are known as kerangas and the poor- side, slightly concave on adaxial side. Adult leaves
est sites may only support heath forest dominated linear-lanceolate, spreading at ca. 45, 24 mm long,
by this conifer and Gymnostoma (Casuarinaceae) 0.80.9 mm wide and 0.20.3 mm thick, straight
which looks in habit like a conifer. In Sabah D. pec or slightly curved forward at apiculate apex, keeled
tinatum may occur on ultrabasic soils supporting abaxially, distinctly concave adaxially. Stomata on
an open, low vegetation of shrubs and ferns, where both sides (leaves amphistomatic) but more abun-
trees remain small. dant on the adaxial side where lines extend to apex.
Pollen cones not observed. Seed cones terminal,
Conservation often on a small lateral shoot, 23 mm long, sub-
tended by reduced leaves less than 2 mm long; cone
Deforestation especially in relation to expanding oil bracts slightly spreading, straight, up to 3 mm long
palm plantations poses a serious threat and has frag- and 0.5 mm wide, covering the basal epimatium of
mented remaining populations. the seed when mature. Seeds solitary or in pairs, at
IUCN: EN [B2ab (iii)] oblique angle to cone or shoot apex, ovoid, slightly
flattened, ca. 4 mm long, brown.
Uses
Taxonomic notes
No uses have been recorded for this species.
De Laubenfels (1969), when first describing this new
species, included collections from Borneo (Sarawak)
in it, but later, in Flora Malesiana (De Laubenfels, Dacrydium suprinii Nimsch, Feddes Repert.
1988) he had concluded that these belong to 118 (12): 52. 2007. Type: New Caledonia: Grande
Dacrydium ericoides (E. F. Brunig SFN 8722, type). Terre, Province Sud, Chtes de la Madeleine, 100
The latter species has longer and slightly wider, m downstream from waterfall, H. Nimsch 5122
straight and distinctly flattened leaves narrowing (holotype B).
abruptly to an apiculate apex. Hence D. spathoides,
as far as is known, is restricted to one locality in New Etymology
Guinea.
This hybrid species was named after Bernard Suprin,
Distribution a French resident botanist with intimate knowledge
370 of the New Caledonian flora.
New Guinea: Papua (Central Highlands, Idenburg
River drainage). Vernacular names
TDWG codes: 43 NWG-IJ
No common names are known for this species.
Ecology
Description
Dacrydium spathoides is known from a single local-
ity on the Idenberg River where it forms a canopy Small shrub-like tree 24 m tall. Branches in mature
tree in moist, mossy montane rain forest at an alti- plants in pseudo-whorls of a few first order branches
tude of 21502200 m a.s.l. only, spreading and ascending to form a candelabra
crown with tufts of foliage branches. Leaves have
Conservation characters intermediate between D. araucarioides
and D. guillauminii; juvenile leaves 1015 mm long,
This species is known from only one locality includ- 0.81 mm wide, adult leaves densely arranged, 510
ing its type specimen, but due to poor collecting mm long, ca. 1 mm wide, rhombic in cross-section,
records from the area nothing can be concluded partly imbricate but free at least towards apex, nar-
from this as to rarity, population size etc. The fact rowly lanceolate with a wide base, gradually taper-
that is has been described in 1969 and no subsequent ing towards the acute-acuminate but non-pungent
collections were made is due to its obscurity as a apex. Stomata few near base on abaxial (lower) face
species, which is not commonly known to botanists and in two lateral lines from base to apex on the
who might visit the area. Rather than maintain it as adaxial side. Pollen cones both terminal and lateral,
DD, the Conifer Specialist Group believes this spe- 1016 mm long and 45 mm wide when expanded;
cies should be flagged as NT as it is surely very rare lateral cones at base of a terminal cone and smaller.
and restricted. As a canopy tree in rain forest it may Microsporophylls with a greatly elongated acicu-
be subject to logging. lar apex 45 mm long near base of cone, gradually
IUCN: NT shortening towards cone apex to 12 mm; pollen sacs
2, partly hidden by curved basal part of microsporo-
Uses phyll. Seed cones terminal on long or short foliage
shoots, ca. 6 mm long, hidden by leaves. Seeds 12
The timber of this tree is possibly used for construc- per cone, proximally covered for half or more by
tion (house building), but its very localized occur- the epimatium and subtended by bracts; bracts leaf-
rence means that no specific data are available. like, long, exceeding the seed; seeds ovoid-oblong,
34 2 mm, with a constricted apex, brown.
Taxonomic notes
Microcachrys W. Archer, Hookers J. Bot. 2: 51. 1850, sionally in whorls of 3, imbricate, scale-like, rhombic,
non Hook. f. (1845). Type not designated. more or less keeled distally, with entire margins and
obtuse apex, 12 0.61.5 mm; stomata in two short
Greek: di = two; selma = upper deck, seat; referring whitish bands in lower part of the leaf, eglandular;
to the fertile scales in the seed cone. young leaves on mature branchlets reddish at first,
turning green or olive-green. Pollen cones numer-
Description ous, terminal and solitary on ultimate branchlets, 373
to 3 mm long; microsporophylls 68, smaller than
See the species description. leaves, triangular-deltoid, usually 4 fertile, each with
2 abaxial pollen sacs. Seed cones terminal on ulti-
Distribution mate branchlets, maturing in one season to small
cones 3 2 mm, consisting of 2 pairs of decussate
As for the species. cone scales surrounding a central columella; upper
scales fertile, 23 11.5 mm, with bract margin vis-
Diselma archeri Hook. f., Fl. Tasmania 1 (5): 353, t. ible at the distal end; columella relatively large, resin-
98. 1857. Type: Australia: Tasmania, Great Western ous-glandular, clavate or ovate-cylindrical, breaking
Tiers, Meander Falls, [summit of the western off in old persistent cones. Seeds 2 (or 1) developing
mountains and Falls of Meander], R. C. Gunn on each fertile bract-scale complex, ovoid-flattened,
[366] s.n. (lectotype K). Fig. 120 brown; wings 2(3), marginal, broad, partly sur-
rounding seed.
Etymology
Taxonomic notes
This species was named after William Archer (1820
1874), who studied the flora of Tasmania. The foliage and leaf arrangement as well as leaf mor-
phology of Diselma archeri very closely resemble
Vernacular names those of the sympatric prostrate shrub Microcachrys
tetragona (Podocarpaceae), which are opposite-
Cheshunt pine decussate (in distichous pairs according to Hill &
Brodribb, 1999), unlike the usually helical arrange-
Description ment in that family. It may be worthwhile to investi-
gate whether the two taxa really have a homologous
Shrubs, or very small trees 1.54 m, occasionally phyllotaxis or whether the podocarpaceous coni-
a small tree up to 6 m in shade, rarely prostrate, fer has in fact a bijugate phyllotaxis (Tomlinson
stem to 3040 cm diam. but mostly smaller, ever- & Zacharias, 2001) as a deviation from a helical
green, dioecious; trunk monopodial, short, or mul- arrangement. It is interesting that the terminal cones
tistemmed, with sprawling or assurgent branches of both sexes in Microcachrys have their sporophylls
from a common base, forming extensive bushes. in a helical arrangement, while those in Diselma are,
Bark scaly, rough, brown turning grey; inner bark like their leaf homologues, decussately arranged.
reddish brown. Branches short, rigid, forming a
dense crown. Foliage branches numerous, not in a Distribution
plane, very dense, straight and more or less rigid,
short (1020 mm), tetragonal or occasionally trigo- W Tasmania, in most highland areas but absent in
nal, ca. 1.5 mm diam., covered with closely appressed the NW.
leaves, persistent. Leaves opposite-decussate, occa- TDWG codes: 50 TAS
Ecology Conservation
This shrubby species occurs in subalpine and alpine This monotypic genus, though endemic to Tasma-
moorland and boulderfields, sometimes ripar- nia, is widely distributed in the highlands, where
ian along streams or on shores of lakes and ponds, almost all populations are reserved within the West-
among perennial herbs and other shrubs (conif- ern Tasmania Wilderness World Heritage Area
erous heath), e.g. Astelia alpina, Pherosphaera (WTWWHA). It is not considered threatened,
hookeriana, Microcachrys tetragona, Podocarpus although it is sensitive to fires, which have in historic
lawrencei, Richea scoparia, R. pandanifolia, Lomatia times increased in frequency in some areas.
spp., Orites acicularis, O. revoluta, Helichrysum spp., IUCN: LC
374 Leptospermum spp. and larger shrubs or stunted
trees, e.g. Athrotaxis cupressoides, Phyllocladus asple Uses
niifolius, Nothofagus gunnii, and Eucalyptus coccifera.
The altitudinal range of Diselma archeri is from 550 This shrub is uncommon in cultivation, but some
m to 1400 m a.s.l. Soils are acidic, often peaty and specialist nurseries grow it for the horticultural
water-logged and shallow over igneous rock. Annual trade. It is also grown in botanic gardens. Diselma
precipitation is very high, snow and frost are pos- archeri (the vernacular name is rarely used) is a
sible year-round, with snow cover at the highest alti- slow growing, more or less erect shrub, depending
tudes lasting several months. on what shoots are being used for propagation by
cuttings, and very suitable for rock gardens. Read
Dwarf is an even slower growing cultivar with light
green young foliage from a plant found on Mt. Read
in the north of Tasmania.
Falcatifolium de Laub., J. Arnold Arbor. 50: 308. 1969. Type: Falcatifolium falciforme
(Parl.) de Laub. [Podocarpus falciformis Parl.] (Podocarpaceae).
Latin: falcis = sickle; folia = leaf. Transitional forms between juvenile and adult leaf
forms also exist and one should consult the descrip-
Description tions before concluding an identification.
Dioecious evergreen shrubs or trees. Resin in bark 1a. Low shrubs. Adult leaves 0.61 cm long, 1.82
and leaves. Bark thin, with scattered lenticels, even- mm wide (only known from one locality in
tually flaking. Branching in monopodial trees in New Guinea) F. sleumeri 375
pseudo-whorls, more irregular in shrubs, foliage 1b. Shrubs or (small) trees. Adult leaves normally
branches terminating in loose buds formed by nar- longer than 1 cm 2
rowly triangular scale leaves. Leaves spirally inserted, 2a. Juvenile leaves narrowly linear-lanceolate, up
dimorphic, more or less appressed scale leaves on to 7 cm long, 1.21.5 mm wide. Trees to 20 m
leading and fertile shoots, alternating with more tall F. angustum
or less distichously spreading, bilaterally flattened, 2b. Juvenile leaves usually shorter than 3 cm, if lon-
obliquely lanceolate-falcate foliage leaves on (lateral) ger (to 12 cm) then wider than 5 mm. Shrubs or
vegetative shoots, latter leaves single-veined, much trees to 22 m tall 3
larger than scale leaves, but variable in size, with sto- 3a. Juvenile leaves nearly linear, 12 cm long, 1.5
mata on both sides (leaves amphistomatic). Pollen mm wide, adult leaves 36 mm wide with
cones on axillary, usually solitary but sometimes obtuse or sometimes acute apex. Receptacle
clustered, short, scaly shoots (peduncles), cylindri- when full-grown 20 8 mm F. taxoides
cal and catkin-like; microsporophylls small, trian- 3b. Juvenile leaves falcate or linear-lanceolate, 2.5
gular or acuminate, with two globose pollen sacs 12 cm long, 3.512 mm wide, adult leaves
containing bisaccate pollen. Seed cones solitary on (1.5)29 mm wide with acuminate or pungent
axillary, scaly dwarf shoots (peduncles), consisting apex. Receptacle when full-grown smaller than
of several spirally arranged bracts; bracts becom- 12 6 mm 4
ing swollen, fleshy and red, forming an irregularly 4a. Juvenile leaves 1012 cm long, 1012 mm wide;
shaped receptacle, with one fertile bract at or near adult leaves falcate or S-curved F. falciforme
apex bearing an inverted ovule on the adaxial side. 4b. Juvenile leaves not longer than 7.5 cm and not
Seeds at an oblique angle to axis of receptacle, erect, wider than 7 mm, adult leaves falcate (some-
surrounded at base by a swollen epimatium, ovoid times at base only), not S-curved 5
but more or less flattened, with two lateral ridges. 5a. Leaves flushing whitish/yellowish green or
glaucous; juvenile leaves only slightly longer
6 species. than adult leaves, to 3 cm long. Pollen cones
0.51.3 cm long F. papuanum
Distribution 5b. Leaves flushing pinkish or purplish; juvenile
leaves markedly longer than adult leaves, to 7.5
Malesia: Peninsular Malaysia, Borneo, Sulawesi, cm long (adult leaves to 4 cm long). Pollen cones
Phulippines, Maluku [Moloccas], New Guinea. SW 56 cm long F. gruezoi
Pacific: New Caledonia.
Falcatifolium angustum de Laub., J. Arnold Arbor.
Key to the species of Falcatifolium 50: 312. 1969. Type: Malaysia: Sarawak, Bintulu, Sg.
Meluang W.S., E. F. W. Brunig S8866 (holotype L).
The reproductive organs (pollen cones and seed
cones) of some species remain not or poorly known; Etymology
for this reason their use in this key remains ten-
tative and should be treated with some caution. The species epithet refers to the narrow leaves.
Vernacular names Conservation
No vernacular names are recorded for this species. Deforestation in relation to the expansion of oil palm
plantations poses a serious threat to this species.
Description IUCN: EN [B1ab (ii, iii, v), B2ab (ii, iii, v)]
Etymology Distribution
This species was named after Hermann Otto Sleumer New Guinea: Birds Head Peninsula (Mt. Nettoti).
(19061993), a German botanist who worked at the TDWG codes: 43 NWG-IJ
Rijksherbarium in Leiden.
380 Ecology
Vernacular names
Stated to occur in dark mossy forest, so presumably
No vernacular name is known for this rare species. a ground covering species under a canopy of trees.
The altitude is given as 1920 m above sea level.
Description
Conservation
Low, more or less flattened or prostrate [?] shrubs
20 cm tall or more [based on a single specimen Known from only one locality and its type speci-
described to be of that dimension this habit and men, but due to poor collecting records from the
size remain speculative]. Bark scaly, greyish brown. area nothing can be concluded from this as to rar-
Branches numerous, spreading to erect, creating a ity, population size etc. It is described as a pros-
spreading, flattened crown covering the ground for trate shrub in dark, mossy forest. The fact that is
several square meters. Leaves linear-ovate, curved at has been described in 1988 and no subsequent col-
the shortly petiolate base, falcate or nearly straight, lections were made is undoubtedly due to its very
0.61 cm long and 1.82 mm wide, midrib raised on local occurrence. Rather than maintain it as DD
the abaxial side; margins slightly revolute, more or the Conifer Specialist Group believes this species
less abrubtly tapering to an apiculate apex with a should be flagged as NT as it is surely very rare and
prickly spine 0.2 mm long. Reproductive structures restricted.
not known. IUCN: NT
From Mt. Nettoti at 1920 m a.s.l. in the Birds Head No uses are recorded of this species; if its habit is
Peninsula a dwarfed plant was collected with very indeed a prostrate shrub it could be of interest to
small leaves 610 2 mm which are morphologi- horticulture, but only in countries with a frost-free
cally similar to leaves on trees of Falcatifolium pap climate.
uanum, in particular sun-exposed leaves. The leaves
were illustrated with a small photograph in the Falcatifolium taxoides (Brongn. & Gris) de Laub.,
protologue of F. papuanum (De Laubenfels, 1969: J. Arnold Arbor. 50: 310. 1969. Dacrydium taxoides
313, fig. 6b) and it was speculated that it might rep- Brongn. & Gris, Ann. Sci. Nat. Bot., sr. 5, 6: 245.
resent a distinct entity or even a distinct species 1866. Type: New Caledonia: Grande Terre, Province
(De Laubenfels, 1988: 374). De Laubenfels & Silba Nord, Balade, [montagnes de Balade], E. Vieillard
(op. cit.) proceeded to describe it as a new species, 1259 [185560] (lectotype P). Fig. 124
F. sleumeri, based on the same specimen already
mentioned in the account of 1969 and with no fur- Etymology
ther knowledge or information, e.g. on reproductive
structures; that single specimen, P. van Royen & H. The species epithet refers to Taxus, presumably from
Sleumer 8203a (holotype L) is apparently sterile.This a resemblance of its leaves with those of yew.
Vernacular names Distribution
No common names are known for this species. New Caledonia: throughout Grande Terre.
TDWG codes: 60 NWC
Description
Ecology
Shrubs or small trees 215 m tall; trunk with max.
d.b.h. 25 cm. Bark smooth becoming rough on larger Falcatifolium taxoides is common as an understorey
stems, brown weathering grey. Crown usually open tree in montane humid forests from lowland hills in
and sparse in sub-canopy trees, more dense in trees the south of Grande Terre to the summit ridges of
growing in open vegetation. Leaves of two types: the highest mountains on the island at 1400 m or 381
scale leaves and foliage leaves. Scale leaves on lead- even more. It occurs on ultramafic soil derived from
ing shoots and on base of lateral shoots, more or serpentine as well as on acidic metamorphic rock
less appressed, subulate to narrowly lanceolate, 23 (micaschist). It is associated with Araucaria mon
1 mm, sometimes transforming into small foli- tana, A. laubenfelsii, A. humboldtensis, and Agathis
age leaves. First leaves on seedlings nearly linear, montana (on Mt. Pani) as well as angiosperm trees
spreading at wide angles, petiolate, 1220 1.5 mm, e.g. in the Meliaceae. Its most peculiar associate is
strongly keeled abaxially, soon replaced by adult the parasitic podocarpaceous conifer Parasitaxus
leaves, flushing leaves pink with whitish bloom, usta, to which it is the only host. This diminutive
becoming bright green on both sides. Adult leaves shrub or treelet usually attaches to its roots and
falcate at base, then more or less straight or curved seems to grow from the litter under Falcatifolium
or ovate-oblong, variable in size on a single branch- taxoides. The large and succulent red receptacles are
let, 0.82.5 cm 36 mm, more or less petiolate at conspicuous and eaten by birds, who disperse the
base, tapering in the distal part; midrib raised on single seed thereby.
both sides or inconspicuous; apex obtuse or some-
times acute. Stomata in numerous intermittent lines Conservation
on both sides of leaves from base to apex. Pollen
cones on axillary or sometimes terminal scaly dwarf IUCN: LC
shoots, often 23 together, 1.52.5 cm 1.52 mm;
microsporophylls terminating in an acuminate apex Uses
above two yellow pollen sacs. Seed cones solitary on
axillary dwarf shoots with acuminate scale leaves, This small tree is not exploited for its timber and
with 1012 scales subtended by elongated bracts; other uses in New Caledonia are unknown. It is
cone swelling to an irregularly shaped receptacle ca. in cultivation only in some collections of botanic
20 8 mm and becoming succulent and bright red; gardens; it could grow outside only in subtropical
mature seed single, ovoid, obliquely attached near to tropical climates. It has attractive pink flushing
apex, 67 34 mm, slightly flattened with two lat- leaves and larger pollen cones and red, succulent
eral ridges, reddish maturing to dark brown. receptacles than other species and would make an
attractive small amenity tree in tropical countries.
Fitzroya Lindl., J. Hort. Soc. London 6: 264. 1851. Type: Fitzroya cupressoides
(Molina) I. M. Johnst. [Pinus cupressoides Molina] (Cupressaceae).
Named after Robert FitzRoy (18051865), who apex obtuse; stomata abaxially in a few lines on
achieved lasting fame as the captain of HMS Beagle decurrent base; leaf colour lustrous green to glaucous
during Charles Darwins famous voyage. green, with whitish wax on stomatal zones. Pollen
cones predominantly terminal, solitary, cylindrical,
Description 68 23 mm; microsporophylls 1524, in alternate
whorls of 3, imbricate, peltate-ovate with acute apex,
382 See the species description. keeled abaxially; margins entire, bearing (2)46
small pollen sacs on abaxial side. Seed cones termi-
Distribution nal or subterminal, formed by 23 whorls of slightly
modified scale leaves, followed by 2 alternate whorls
As for the species. of 3 fertile scales, maturing in 1 season to cones with
wide spreading scales 1014 mm diam. Bract-scale
Fitzroya cupressoides (Molina) I. M. Johnst., Contr. complexes thin woody, obovoid-spathulate, largest
Gray Herb., n. s. 70: 91. 1924. Pinus cupressoides one ca. 5 4 mm, with thickened, wrinkled apical
Molina, Sag. Stor. Nat. Chili: 168. 1782. Type: Chile: part and subapical, curved umbo of exserting bract
Los Lagos, Cordillera de la Costa, 30 km from San apex 12 mm long. Columella variably shaped, trigo-
Juan de la Costa, J. L. Morrison 17636 (neotype K). nal to tripartite, ca. 4 mm long, flattened, not woody,
Fig. 125, 126 resinous-glandular, more or less translucent. Seeds
usually only well developed on upper fertile whorl,
Etymology 34 2 mm, ovoid-oblong, flattened, with a con-
cave hilum; wings usually 2, sometimes 3, crescent-
The species epithet means like a cypress shaped with undulate margins, ca. 2 mm wide.
(Cupressus).
Distribution
Vernacular names
S Argentina: Chubut, Neuqun, Rio Negro; S Chile:
Patagonian cypress; Alerce (Spanish), Lahun Los Lagos. For the most part, the still existing popu-
lations of Fitzroya cupressoides are in Chile, now dis-
Description junct in the coastal Cordillera, some in the Central
Valley, and others in the Andes, where they cross the
Trees to 50(60) m tall, evergreen, dioecious, rarely border into Argentina.
monoecious; trunk monopodial, straight, buttressed TDWG codes: 85 AGS-CB AGS-NE AGS-RN CLS-LL
in old individuals and often with root suckers, to
35 m diam. above buttress. Bark on trunk thick, Ecology
with deep fissures, exfoliating in long strips, reddish
brown. Branches spreading horizontally or curved A dominant or emergent tree usually forming
down, forming a conical or pyramidal crown. Foliage more or less large stands (alerzales) scattered in
branches spreading or pendulous, ultimate branch- the Valdivian evergreen rainforest. Understorey
lets 14 cm long, more or less terete, densely covered tree species are e.g. Podocarpus nubigenus, P. salig
with leaves, persistent. Leaves scale-like, in alternate nus, Prumnopitys andina, Saxegothaea conspicua,
near-whorls of 3, imbricate, decurrent, with distal Pilgerodendron uviferum, Nothofagus betuloides, N.
part variously (re)curved; apex incurved but free, dombeyi, N. nitida, N. obliqua, Drimys winteri, and
lanceolate to ovate, 46 mm long on younger trees, Laurelia philippiana. There are various forest types
23 mm long on older trees, 12 mm wide, keeled with shifting dominance of these associates on altitu-
abaxially, adaxially with a midrib; margins entire; dinal gradients and on soil types, with e.g. a very wet,
acidic woodland bog type at middle to high (1200 m) The most notable private property where impor-
elevation dominated by Pilgerodendron and Fitzroya tant populations of Fitzroya are protected is The
and a shrublayer of the conifer Lepidothamnus fonkii Pumalin Park in southern Chile which was set up
in carpets of Sphagnum. Another forest type occurs in 1991 to help protect 300,000 ha of pristine rain-
between 8001200 m on better drained volcanic forest, making the largest private park in the world.
soils with dominance of Nothofagus betuloides and In the late 1990s, studies carried out into the genetic
emergent Fitzroya; it is there dependent on episo- variability of Fitzroya using DNA markers have
dal disturbance in the absence of which Nothofagus helped guide researchers from the Forestry Faculty
would prevent its regeneration. Fitzroya cupressoi of the Universidad Austral de Chile in establishing
des is one of the longest living trees in the world, an important restoration programme in the Central
with record ages counted between 3500 and 4000 Depression. 383
years. The soils on which Fitzroya occurs are usually IUCN: EN (A2cd)
poorly drained and water-logged, or incipient soils
on volcanic ash deposits, at lower elevations sandy Uses
or loamy soils predominate, often with a very high
organic content and a low pH (4.05.1) in the upper This species has been extensively used for build-
layer. Mean annual precipitation ranges from 2000 ing timber and especially shingles for roofing. Its
6000 mm, mostly as winter rains, with snow at the extremely rot resistant wood is so durable, that
higher elevations. shingles on roofs 130150 years old are still perfectly
sound (Golte, 1996). In 1973 logging was still going
Conservation on and sawmills were well stocked and busy, but
exploitation has declined sharply since about 1980,
The range of Fitzroya cupressoides has historically with export no longer legally possible (an illegal
been reduced greatly by centuries of logging, prob- shipment of wood was intercepted in the UK and
ably aggravated by recurring fires both natural and given to the Royal Botanic Gardens, Kew, who built
man-made. It has largely disappeared from the low- an education centre from it at its garden Wakehurst
land areas N of Puerto Montt, where it was once Place in Sussex, England). This conifer was intro-
extensive and abundant (Golte, 1996). It is not doing duced to cultivation by William Lobb in 1847 (possi-
well in many of the remaining, mostly upland and bly based on his brothers collection) and it remains
very wet stands, and large tracts of land merely have an uncommon tree mostly confined to dendrologi-
standing dead snags with little or no natural regen- cal collections. Recently renewed seed collecting
eration. Exploitation has been on a devastating scale has replenished the original introduction, which
with no intent of sustainability until 1976, when log- appeared to have been propagated from cuttings;
ging of living trees was banned by the Chilean gov- new plants have been distributed in the UK largely
ernment and the species was declared a National as ex situ conservation stock by the International
Monument. Fires continue to kill trees, which can Conifer Conservation Programme of the Royal
then be logged and the land converted to pasture. Botanic Garden at Edinburgh, Scotland. As a CITES
Since 1973 the species is listed on CITES Appendix Appendix I listed species its commercial distribution
I which prohibits all export. Although it is afforded will remain restricted and plants can only be sold if
protection in both Argentina and Chile in National they originate from legal stock in cultivation outside
Parks, most populations occur on private property. Chile and Argentina.
Fokienia A. Henry & H. H. Thomas, Gard. Chron., ser. 3, 49: 67. 1911. Type:
Fokienia hodginsii (Dunn) A. Henry & H. H. Thomas [Cupressus hodginsii Dunn]
(Cupressaceae).
Fokien is an earlier European spelling for Fujian, a on flattened ultimate branchlets but increasingly less
province of China. so on older (whip) shoots, becoming long decur-
rent to 15mm, with facials usually smaller and/or
Description narrower than the laterals, green or glaucous green.
Facial leaves appressed, obtuse, variable in size from
384 See the species description. 27 12mm on ultimate branchlets; stomata in
two bands separated by a midrib. Lateral leaves lon-
Distribution ger than facials or nearly equal in length, bilaterally
flattened, variable, 410 23(4) mm on ultimate
As for the species. branchlets; apex incurved, free to appressed, acute to
obtuse; stomata in a single whitish band, eglandular.
Pollen cones terminal, 45 1.52mm, initially sub-
Fokienia hodginsii (Dunn) A. Henry & globose but elongating when maturing, with 1012
H. H. Thomas, Gard. Chron., ser. 3, 49: 6668, decussate, broadly ovate-acute microsporophylls
f. 3235. 1911. Cupressus hodginsii Dunn, J. Linn. each bearing 3 small, abaxial pollen sacs. Seed cones
Soc., Bot. 38: 367. 1908; Chamaecyparis hodginsii terminal, maturing in the second year, caducous, to
(Dunn) Rushforth, J. Biol. (Viet Nam) 29 (3): 38. 1525 1222mm, subglobose, brown. Bract-scale
2007. Type: China: Fujian, Min Jiang, Nanping, complexes (10)1216, decussate, spreading at matu-
[Yenping], A. E. N. Hodgins HK 3505 (holotype rity, to 10 16mm (uppermost 24 much smaller and
K). Pl. 14, Fig. 127 sterile), cuneate-peltate, flattened, woody, with irreg-
ular margins, distally with a central depression and a
Etymology small mucronate umbo (bract tip), dull brown, with
striated anterior part, orange- to reddish brown with
The species epithet is after A. E. N. Hodgins, who whitish seed marks. Seeds 2030 per cone, 45mm
collected it in 1905. long, ovoid and 34 ridged, with acute apex and trun-
cate base (hilum), brown, wings 2, very unequal in
Vernacular names size and shape; larger wing ca. 6 5mm, smaller one
ca. 1.5mm, or a mere strip near the seed apex.
Fujian Cypress; fu jian bai (Chinese)
Taxonomic notes
Description
Some authors have placed this species either in
Trees to 30 m tall, evergreen, monoecious; trunk Cupressus or (recently) in Chamaecyparis based on
monopodial, to 1 m d.b.h. Bark on trunk nearly comparisons of (some) gross-morphological char-
smooth, becoming irregularly fissured and flak- acters. Detailed analyses of the ontogeny of the seed
ing, purplish brown. Branches long, spreading or cones (Jagel, 2001; Farjon, 2005a) and phylogenetic
down-curved, self-pruning lower down the trunk, analyses based on morphology (Farjon, 2005a) as
forming a broadly pyramidal, in old trees rounded well as DNA sequence data (Gadek & Quinn, 1993)
and dense crown. Foliage branches spreading in flat- have demonstrated that Fokienia hodginsii represents
tened sprays (plagiotropic), ultimate branchlets pin- an evolutionary lineage separate from these two gen-
nately arranged, forming frond-like sprays, but in era and therefore merits recognition at the rank of
older trees of irregular and shorter length forming genus. Similarities observed, especially in the foliage
flat, rounded sprays, deciduous. Leaves decussate, and in later (mature) phases of cone development,
imbricate, decurrent, scale-like, strongly dimorphic are likely to be the result of convergent evolution.
385
plate 14. Fokienia hodginsii. 1. Habit of tree. 2. Branch with foliage. 3. Branchlet with juvenile leaves
(lower side). 4. Branchlet with adult leaves. 5. Pollen cones. 6. Microsporophyll with pollen sacs. 7. Seed
cones. 8, 9. Seeds.
Distribution using helicopters and possibly beyond sustainabil-
ity, because the dark red timber is considered very
China: Chongqing, Fujian, N Guangdong, Guangxi, valuable (roughly estimated at US$ 2500/m3). The
Guizhou, S Hunan, W Jiangxi, SE Sichuan, SE situation in China is possibly similar in many areas
Yunnan, S Zhejiang; N Lao PDR; Viet Nam. where mature forests are shrinking under popula-
TDWG codes: 36 CHC-CQ CHC-GZ CHC-SC tion pressures. However, the species is very widely
CHC-YN CHS-FJ CHS-GD CHS-GX CHS-HN CHS-JX distributed and regeneration has been observed in
CHS-ZJ 41 LAO VIE most populations. Continued logging at current
or increased rates may cause scarcity of large trees
Ecology in the near future and the logging ban in China, if
386 effective, does not apply to other countries.
This conifer is usually a minor constituent of sub- IUCN: VU [A2acd; B2ab (iiv)]
tropical to warm temperate evergreen (mixed)
mesophytic forest, which in undisturbed state is Uses
dominated by numerous angiosperm trees e.g.
Castanopsis spp., Quercus spp., Lithocarpus spp., The wood of this species, like other cupressaceous
Nyssa sinensis, Pasania spp., Schima argentea, S. trees in eastern Asia, is much valued for construc-
superba, and a few other conifers e.g. Cephalotaxus tion. In most areas where it occurs, wild grow-
fortunei and Nothotsuga longibracteata. With contin- ing trees are logged; its slow growth makes it a
ued disturbance conifers like Cunninghamia lanceo- tree difficult to exploit economically in plantation
lata and especially Pinus massoniana or P. densata forestry. In southern China and in Viet Nam the
become more abundant and eventually dominate. rot resistant and even-grained wood is prized for
The altitudinal range of Fokienia hodginsii is from doors, window frames, and carved panels, as well
350 m to 2100 m a.s.l. Soils are acidic (pH 56) yel- as furniture. In remote regions pit-sawn timbers
low or brown loamy sands over sandstone, shales, are often transported off the mountains by man-
granite, or rhyolite and well drained. Mean annual power to fetch high prizes with carpenters in the
precipitation is 1200mm or more. towns. This conifer is cultivated in cemeteries and
parks in China, but rare elsewhere despite its rela-
Conservation tive hardiness and suitability in parts of the world
with similar climates such as New Zealand, Atlantic
Mature trees are selectively logged in Viet Nam and SW Europe, northern California and Oregon and
Lao PDR, even from forest reserves in remote areas southern Japan.
Glyptostrobus Endl., Syn. Conif.: 69. 1847. Type: Glyptostrobus pensilis (Staunton ex
D. Don) K. Koch [Thuja pensilis Staunton ex D. Don] (Cupressaceae).
Greek: glypto- = cut into; strobus = cone; referring to with leaves. Leaves alternate to helically arranged,
the margins of the seed cone scales. of three types, all green with white stomatal dots.
Scale leaves on long shoots decurrent, imbricate,
Description 1.53 0.50.8mm, lanceolate, convex or keeled
abaxially; apex incurved but free, obtuse. Needle-
See the species description. like leaves on short shoots of two types, helically
arranged, decurrent, and either more or less disti- 387
Distribution chous, linear, 1530 1.53mm, bilaterally flattened,
with entire margins, acute, or more commonly in
As for the species. ranks of 3 or nearly radially spreading, subulate,
410 0.40.6mm, abaxially keeled and adaxially
grooved. Stomata on all leaves types on both sides,
Glyptostrobus pensilis (Staunton ex D. Don) but more on adaxial side. Pollen cones terminal,
K. Koch, Dendrol. 2 (2): 191. 1873. Thuja pensilis solitary, 35mm long, subglobose to ellipsoid, with
Staunton ex D. Don, in Lambert, Descr. Pinus, 1520 helically arranged, imbricate, ovate-convex
ed. 2, 2: 115. 1828. Type: China: Guangdong, microsporophylls bearing (2)46(9) abaxial,
[Province of Guangton], G. L. Staunton s.n. small pollen sacs. Seed cones terminal, solitary on
(holotype BM). Fig. 128 determinate branchlets, subtended by a few short,
broad, keeled leaves grading into bract-scale com-
Etymology plexes, maturing in two seasons; mature cones
pyriform to obovate, 1425 1015mm, with 2025
The etymology of the species epithet is unclear, helically arranged, imbricate woody scales, matur-
perhaps it derives from Latin penicillatus = pencil- ing brown. Bract-scale complexes increasing in size
shaped [the branchlets]. to 13 6.5mm, obovate-oblong, cuneate proximally,
rounded distally, connate but parting slightly at
Vernacular names maturity, with 48(11) subapical tooth-like lobes ca.
1mm long (often worn off) along distal margin and
Chinese Swamp Cypress, Chinese Water Fir; shui a (sub)central, acute bract apex 23 23mm; abax-
song (Chinese) ial surface grooved or rough; adaxial surface with
two unequal, deep seed cavities. Ovules 2 in axil of a
Description fertile bract, erect, in well developed cones maturing
to 2030 seeds; seeds elliptic, 57 34mm, slightly
Trees to 15(25) m tall (one planted tree in China flattened, often grooved, brown, with a single proxi-
is 39 m tall), semi-evergreen, monoecious; trunk mal, oblong and thin wing of 47 24mm.
markedly buttressed at base especially when grow-
ing in watery substrate, to 1.21.5 m d.b.h. Bark on Taxonomic notes
trunk thin with long, irregular strips, whitish brown
or grey-brown. Branches long, slender, spread- In many descriptive works on conifers, the name
ing horizontally or curved down in old trees, foli- Glyptostrobus lineatus (Poir.) Druce has been applied
age in distinct tufts at ends of branches, forming a to this taxon. That combination by Druce, published
pyramidal or more rounded, open crown. Foliage in 1916, was based on Thuja lineata Poir., a species
branches alternate, spreading at less than 45 degrees, newly described in Supplement Vol. 5 of Lamarcks
variable in length, dimorphic, long shoots persis- famous Encyclopdie methodique (Botanique)
tent, slender and flexible, short shoots from termi- in 1817. This description was based on a cultivated
nal or dormant lateral buds, deciduous, all covered plant in the garden of Monsieur Noisette undoubt-
edly in France. Needless to say, this plant cannot to the subtropical to tropical coastal lowlands, but
be traced and there is no specimen in the Paris avoids saline and alkaloid soils or water. A probably
Herbarium (P) that can be linked to it. The descrip- indigenous occurrence in Viet Nam along a marshy
tion fits Taxodium distichum var. imbricatum and small river at greater altitude (around 700 m) could
not Glyptostrobus pensilis and while the former had mean that it also occurs along inland waters. This
been introduced to Europe and taken into cultiva- species was widespread in the Tertiary and may well
tion for more than two hundred years in 1817, the have occurred more widely along upland streams.
latter has been introduced first in England about a
hundred years ago, but rarely endured our climate... Conservation
and seems to be unknown in the living state on the
388 continent (Henry & McIntyre, 1926). The plant in The present natural distribution of this species in
Mr Noisettes garden could not have been a speci- China is possibly restricted to several isolated locali-
men of Glyptostrobus. ties along upland streams in three or four provinces
of SE China. However, it has been widely planted
Distribution along water courses in lowland S China for a very
long time and it may now be impossible to distin-
SE China: Fujian, Guangdong; possibly still natural guish between regional introductions and origi-
in SE Yunnan; extinct in the wild in N Viet Nam. nal occurrence, as many trees are found in ancient
This species has also been found in S Viet Nam cultivated landscapes. In N Viet Nam it formerly
(Gaussen, 1967; Hiep & Vidal, 1996). The collections occurred along the Hong (Red) River but no natu-
there cited from Dac Lac: Buon Uik and Krong Buc ral stands are to be found there at present. Habitat
are said to have been found at ca. 500700 m a.s.l. change due to intensive agriculture is the main cause
and to come from trees growing in swampy forest of its decline. In S Viet Nam two small populations
or in basaltic terrain.; its indigenous status in S Viet are probably indigenous and its status there is CR.
Nam has recently been verified ecologically (Hiep IUCN: CR [C2a(i)]
et al., 2004). Gaussen (1967) wrote that it was already
found in 1955 dans les marcages de B. Vit 60 km Uses
au NE de Ban Mthut, province de Darlac. Several
localities in China that are situated near streams at The rather soft, yellowish wood is like most cupres-
altitudes from 300 to 1000 m a.s.l. away from the saceous wood decay resistant and finds uses in China
coastal plain may well have natural populations, too, ranging from furniture to building of bridges. The
but this needs to be veryfied. Recently, it was also wood of the roots is very light and due to its buoy-
discovered on the Nakai Plateau in Khammouan ancy it is used in China to make life-saving rings. This
Province, Lao PDR, where is now known from 56 species is widely cultivated in southern China and
tiny sub-populations (Thomas & Le Page, 2011). planted along rivers and canals as well as in parks;
TDWG codes: 36 CHS-FJ CHS-GD 41 LAO VIE except for the latter localities mostly to harvest the
timber followed by replanting. This conifer is a truly
Ecology semi-aquatic tree requiring permanent water logged
soil to thrive; planting it in rather than adjacent to a
On river floodplains and in deltas, always near or in pond enhances its chances of survival. It can easily
water, where it develops a buttressed base and occa- survive mild frosts in this habitat. It can be grafted
sionally pneumatophores; also extensively planted onto rootstock of Taxodium, another conifer that
along rivers and canals. A heliophilous species, grows well standing in water (but equally outside it).
intolerant of competition and usually growing in Its autumnal foliage colour is a deeper reddish than
pure stands or solitary along streams. It is restricted that of Taxodium and it retains its leaves longer.
Halocarpus Quinn, Austral. J. Bot. 30 (3): 317. 1982. Type: Halocarpus bidwillii
(Hook. f. ex Kirk) Quinn [Dacrydium bidwillii Hook. f. ex Kirk] (Podocarpaceae).
Greek: halos = the sea; karpos = fruit. 2a. Erect, dense shrubs or small trees to 10 m tall.
Juvenile leaves (5)818 mm long, 11.5 mm
Description wide. Branchlets with adult scale leaves 23mm
thick, angular H. biformis
Dioecious, evergreen shrubs or trees. Resin ducts (1) 2b. Trees to 25 m tall. Juvenile leaves 2045 mm
in leaves; resin in bark and leaves. Bark thin, hard, long, 24mm wide. Branchlets with adult scale
with or without lenticels, becoming scaly and exfo- leaves 1.53mm thick, nearly terete 389
liating in thin flakes. Branching variable, in shrubs H. kirkii
often layering, irregular and ascending in trees.
Terminal buds on shoots with juvenile leaves only.
Leaves spirally inserted, dimorphic, radially spread- Halocarpus bidwillii (Hook. f. ex Kirk) Quinn,
ing lanceolate-linear or linear leaves alternating with Austral. J. Bot. 30 (3): 317. 1982. Dacrydium bid-
appressed, rhombic scale leaves, amphistomatic. willii Hook. f. ex Kirk, Trans. New Zealand Inst.
Pollen cones solitary or with 23 together at apex 10: 388. 1878. Type: New Zealand: South Island,
of scale-leaved branchlets, small, with a few spirally Nelson, J. C. Bidwill 130 (lectotype K, sheet 1, here
arranged, reddish microsporophylls and two glo- designated). Fig. 129, 130
bose, yellow pollen sacs containing bisaccate pollen.
Seed cones solitary at apex of scale-leaved branch- Etymology
lets, consisting of a few slightly enlarged, spreading
bracts continuing from and similar to the scale leaves The species epithet commemorates John Carne
and distinguished by a reddish colour from normal Bidwill (18151853) an early collector of plants in
leaves; one or more fertile with 1 inverted ovule on New Zealand.
the adaxial side. Seeds 15 per cone, becoming erect,
at base surrounded by the swollen epimatium form- Vernacular names
ing a white or yellow collar; seed mostly protruding,
more or less ovoid but flattened, with two lateral Bog pine, Mountain pine, Tarwood
ridges, striated, maturing to lustrous black.
Description
3 species.
Prostrate shrubs, densely branched erect shrubs,
Distribution or sometimes a dwarfed tree to 4 m tall and 35cm
stem diam., commonly forming extensive thickets
New Zealand. by layering. Bark becoming rough, hard and greying
with age. Leaves of two distinct types: juvenile and
Key to the species of Halocarpus adult. Juvenile leaves on seedlings and young plants,
spreading radially, linear, (3)510mm long, 0.8
1a. Prostrate to erect shrubs, sometimes a dwarfed 1.4mm wide, convex adaxially, concave abaxially,
tree to 4 m tall. Juvenile leaves on seedlings and with a midrib on that side; apex obtuse. Adult leaves
young plants. Swollen epimatium at the base of abruptly appearing above juvenile leaves, scale-
the seed white H. bidwillii like, arranged in high spirals, on ultimate branch-
1b. Erect shrubs or trees to 25 m tall. Juvenile lets seemingly decussate, completely clasping the
leaves on seedlings, shrubs and trees. Swollen branchlets (including leaves 1.52mm thick), rhom-
epimatium at the base of the seed yellow or bic in appearance, 1.5 1.2mm (larger on thicker
orange 2 branches), weakly keeled and obtuse. Stomata of
juvenile leaves on both sides (leaves amphistomatic) 1500 m a.s.l. but in the far south on Stewart Island
but in more conspicuous bands on either side of the it comes down to near sea level. It can be associated
midrib on the adaxial side, on adult leaves scattered with H. biformis, Podocarpus nivalis, Phyllocladus
to just below apex. Pollen cones often numerous, ter- trichomanoides var. alpinus, Gaultheria depressa,
minal on branchlets with adult leaves, 35mm long, and Cassinia vauvilliersii in the central part of North
slightly wider than branchlet; microsporophylls Island in similar open habitats, or with podocarp
similar to scale leaves, yellow with 24 red pollen trees like Prumnopitys taxifolia, P. ferruginea and
sacs. Seed cones subterminal and erect on branch- Manoao colensoi in open coniferous forest at lower
lets with adult leaves, solitary, consisting of 45 leaf- altitudes further south.
like, slightly swollen bracts; one or sometimes two
390 bracts fertile; epimatium aril-like, swollen and fleshy Conservation
with irregular but clefted upper margin, white, sub-
tending and partially enclosing seed; seed 34mm IUCN: LC
long, striated and crested, ripening from green
to black. Uses
Ecology This species was named after the New Zealand bota-
nist Thomas Kirk (18281898), who studied the for-
Halocarpus biformis occurs more often in forested est flora of that country.
habitats than H. bidwillii, but in boggy terrain in
the central volcanic plateau of North Island the two Vernacular names
may occur together. The altitudinal range of this
species is between 550 m to 1400 m a.s.l. As a small monoao, manoao (Maori)
Description forest to a maximum altitude of ca. 700 m. In the
Waitakere Ranges near Auckland the kauris (Agathis
Trees to 25 m tall; trunk to 1 m d.b.h., short with australis) form emergent trees, with Dacrycarpus
large, ascending branches forming a rounded or flat- dacrydioides, Dacrydium cupressinum, Podocarpus
topped crown. Bark smooth with horizontal lenticels cunninghamii, Prumnopitys ferruginea, and oca-
on young trees, becoming hard, scaly and weather- sionally Halocarpus kirkii together with various
ing grey, with exfoliating flakes covered in small pus- angiosperms making up the main canopy below
tules exposing reddish brown bark. Leaves of two them. Tall tree ferns (Cyathea, Dicksonia) often are
distinct types: juvenile and adult. Juvenile leaves on abundant and can reach the canopy as well. These
seedlings to trees up to 10 m tall, spreading radially, are rainforests with abundant epiphytes in the larger
392 linear, 2045mm long, 24mm wide, petiolate and trees and numerous smaller trees and shrubs in the
twisted at base, flat adaxially, with slightly revolute understorey.
margins abaxially, with a faint midrib on either side;
apex obtuse. Adult leaves abruptly appearing above Conservation
juvenile leaves or on separate branches, scale-like,
arranged in high spirals and on ultimate branchlets This is the only species in the genus to attain large
seemingly decussate, completely clasping branchlets tree size and consequently it has been subject to
(including leaves 1.53mm thick and nearly terete), overexploitation, with many of the forests in which
rhombic in appearance, 1.21.5 1.5mm (larger it occurred now gone. It has a limited natural dis-
on thicker branches), strongly keeled and obtuse. tribution and populations are now fewer and more
Stomata of juvenile leaves on both sides (leaves scattered than before logging began in the 19th
amphistomatic) but more densely in bands on either century. The reduction that occurred is difficult to
side of midrib on adaxial side, on adult leaves con- estimate, but is likely to have been in excess of 50%
spicuous, scattered to just below apex. Pollen cones from its original area of occupancy (AOO), while
often numerous, solitary or 23 together, terminal on podocarp wood was being logged preferentially
branchlets with adult leaves, 34mm long, slightly where forests were not removed altogether. The
wider than branchlet; microsporophylls similar to decline has slowed and ceased in the second half of
scale leaves, green turning yellow, with 46 reddish the twentieth century. It is considered a relatively
pollen sacs. Seed cones subterminal and erect on uncommon tree even in forest reserves that have not
branchlets with adult leaves tinged with red below or only lightly been logged. Regeneration is prob-
cones, solitary, consisting of 45 leaf-like, slightly ably dependent on episodal disturbance of the for-
swollen bracts; one or sometimes two or three bracts est canopy and may seem poor over shorter time
fertile; epimatium aril-like, swollen and fleshy with intervals.
irregular but clefted upper margin, orange, subtend- IUCN: VU A1 (a, c, d)
ing and partially enclosing a seed; seed 34mm long,
striated and crested, ripening from green to black. Uses
Large shrub to 5 m, or small tree to 10 m, dioe- This taxon, formerly recognized as a variety under
cious, rarely monoecious; multistemmed or with Juniperus coahuilensis (see e.g. Adams, 2004, Farjon,
a short trunk and a maximal diam. of 50cm. Bark 2005a), has been found in a recent phylogenetic
of thick branches and trunk fibrous, exfoliating analysis based on DNA sequence data of one-seeded,
in strips, weathering ash-grey. Branches numer- serrate-leaf junipers of the SW USA and north-
ous, long, ascending from base or spreading, often ern Mexico to be quite unrelated to that species. It
curved, those of higher orders ascending or spread- appears to be forming a clade with J. occidentalis
ing, forming a broad, rounded or more irregular and and J. osteosperma, whereas J. coahuilensis is nearer
open crown. Foliage branches numerous, irregularly J. monosperma and J. pinchotii. Therefore, Adams
disposed but not pendulous, ultimate branchlets elevated it to species rank, which on the given evi-
spreading 3070 degrees, slender and stiff, up to dence seems to be convincing and is here followed.
Its morphology is, however, most similar to that of J. brown weathering grey. Branches long, spreading
coahuilensis and the two may not be so easy to sepa- to ascending or nearly erect, forming a sympodial,
rate in the field, as the only reported consistent dif- broad, rounded and dense or more irregular and
ference is in the size of the leaf glands. open crown. Foliage branches numerous, irregularly
disposed but not pendulous, ultimate branchlets
Distribution spreading to erect, stiff, 510(20) 0.91.3mm,
mostly 4-sided in cross section, covered with closely
SW USA: Arizona, SW New Mexico; Mexico: NE appressed leaves, persistent. Leaves decussate, or
Sonora. occasionally in alternate whorls of 3 on whip shoots,
TDWG codes: 76 ARI 77 NWM 79 MXN-SO (slightly) imbricate, scale-like, 12 0.81.2mm
on lateral branchlets, decurrent, rhombic, often 399
Ecology keeled, acute; margins minutely denticulate; sto-
mata on the abaxial side limited to decurrent leaf
In Bouteloua grassland and on adjacent rocky slopes. base, on the adaxial surface in two bands; glands
It has a capacity to coppice, considered weedy in obscure or conspicuous on whip shoot leaves, hemi-
some areas by sheepmen and cattlemen. In Arizona it spheric or oval to elongate, raised; exudate absent;
occurs with Juniperus osteosperma in some localities; leaf colour light to dark green. Pollen cones numer-
other associated species are sometimes Opuntia spp. ous, terminal, solitary, subglobose to ovoid, 24
or Yucca spp., but commonly its only associates are 2mm; microsporophylls 610, decussate, peltate
grasses. Altitudinal range 9801600(2200) m a.s.l. with minutely erose-denticulate or entire mar-
gins and with 34(5) abaxial, large pollen sacs.
Conservation Seed cones terminal on straight short branchlets,
maturing to dark blue within 1 year, globose to
IUCN: LC broadly ovoid, (5)69mm, succulent and resin-
ous. Bract-scale complexes entirely fused, 46,
Uses decussate, indiscernible in mature cones; surface
smooth, rugose when dry, lustrous or very glaucous
No uses have been recorded of this species. (in some populations brownish). Seeds 1(2, rarely
3) per cone, broad ovoid, 46 34.5mm, not flat-
tened, smooth or vaguely grooved, lustrous yellow-
Juniperus ashei J. T. Buchholz, Bot. Gaz. ish brown to chestnut brown, with a lighter hilum
(Crawfordsville) 90 (3): 329. 1930. proximally.
Etymology Distribution
The species epithet commemorates William W. Central USA: Arkansas, Missouri (Ozark Mts.),
Ashe, who made the first botanical collections of it. Oklahoma, Texas (Edwards Plateau, W Texas);
Mexico: N Coahuila.
Vernacular names TDWG codes: 74 MSO OKL 77 TEX 78 ARK 79
MXE-CO
Ashe juniper, Mountain cedar, Ozark white cedar,
Rock cedar, Post cedar, Mexican juniper Ecology
Uses
Juniperus barbadensis L., Sp. Pl. 2: 1039. 1753.
In the past the wood was used by cattle ranchers for
fenceposts, but metal has displaced it. The wood is now Etymology
mainly used to produce cedarwood oil through steam
distillation. This species is not known to be in cultiva- The species epithet refers to the island of Barbados.
400 tion, but it may be present in a few botanic gardens.
Vernacular names
2 varieties are recognized:
West Indies juniper; Barbados cedar (var. barbaden-
sis), Red cedar; Cdre (French) (var. lucayana)
Juniperus ashei J. T. Buchholz var. ashei. Sabina
ashei (J. T. Buchholz) A. V. Bobrov & Melikyan, Description
Komarovia 4: 81. 2006. Type not designated.
Shrubs or trees to 1012 m tall; trunk to 40cm
Description diam., dioecious. Bark on tree trunks thin, stringy,
exfoliating in long strips or thin plates, becoming
Glands on whip shoots hemispheric. Seed cones grey-brown. Branches spreading horizontally or
79mm diam.; seeds mostly 1 per cone, rarely 2. ascending, in shrubs ascending or erect, forming
initially pyramidal, then open and irregular crowns
Distribution in trees. Foliage branchlets numerous, irregularly
disposed or sometimes more or less distichous and
Central USA: Arkansas, Missouri (Ozark Mts.), gradually shorter, variable in length from 530(60)
Oklahoma, Texas (Edwards Plateau). mm, slender or very slender, 0.81.2mm diam.,
TDWG codes: 74 MSO OKL 77 TEX 78 ARK subterete to quadrangular in branchlets with scale
leaves, persistent. Leaves decussate, decurrent,
Conservation scale-like on mature plants and branches, imbricate,
appressed but slightly diverging, on ultimate branch-
IUCN: LC lets 11.5mm long, rhombic to rhombic-lanceolate;
margins entire; apex obtuse to acute or mucronate,
abaxially with a more or less conspicuous rounded
Juniperus ashei J. T. Buchholz var. ovata to elliptic gland, green or greyish green; stomata few
R. P. Adams, Phytologia 89 (1): 17. 2007. Type: at base abaxially, in one or two concave fields adaxi-
USA: Texas, Crockett Co., 5 km W of Ozona, ally. Pollen cones terminal, oblong, 2.53 1.5mm,
R. P. Adams 7463 (holotype BAYLU). more or less quadrangular; microsporophylls 1216,
decussate, broad peltate, with rounded and entire
Description upper margin, abaxially bearing 23 globose pollen
sacs. Seed cones terminal on erect, short ultimate
Glands on whip shoots oval to elongate. Seed cones branchlets, maturing in one season, becoming irreg-
(5)6(8) mm diam.; seeds more often 2 instead of ularly globose or broad pyriform to nearly reniform,
1 per cone. 45 67(8) mm, usually soft pulpy, resinous, pru-
inose-blue or blackish blue. Bract-scale complexes
Distribution 4, decussate, entirely fused, sutures or bract tips
invisible. Seeds (1)23(4) per cone, ovoid-globose
USA: W Texas; Mexico: extreme N Coahuila. (if more than two in a cone flattened on one side),
TDWG codes: 77 TEX 79 MXE-CO 23mm, light yellowish brown.
Distribution Conservation
Bahamas, Cuba, Haiti, Jamaica, Windward Islands. The population that apparently once existed on
TDWG codes: 81 BAH CUB HAI-HA JAM WIN-SL Barbados disappeared probably in the early part of
the 18th century, the location having been converted
Ecology to sugar cane plantations. Recently this variety has
been found on the Petit Piton summit on St. Lucia
In shrubland and in pine barrens or pine savannas, at ca. 730 m a.s.l., where Adams (1989) found ca. 25
in the latter associated with Pinus caribaea var. baha- trees, all within 30 m of the rocky summit, where
memsis, P. tropicalis, in upland hills with P. caribaea they seem to have escaped attention from goats as
var. caribaea or P. occidentalis, also scattered on well as people. Despite the possibility of its occur- 401
rocky cliffs and escarpments in nutrient poor sand or rence on other nearby islands, nothing has been
rocks, commonly over limestone or karst. In the pine found there to date, and this appears to be the only
savannas fires are very frequent. Its altitudinal range extant population of this variety.
is from 8 m to 1600 m a.s.l. The climate is tropical, IUCN: CR (D)
with a mean annual temperature above 20 Celsius
and annual precipitation ca. 10001800mm with a
prolonged dry season. Juniperus barbadensis L. var. lucayana (Britton)
R. P. Adams, Phytologia 78: 145. 1995. Juniperus
2 varieties are recognized: lucayana Britton, N. Amer. Trees: 121. 1908.
Type: Bahamas: New Providence, Southwest Bay,
N. L. Britton & L. J. K. Brace 497 (holotype NY).
Juniperus barbadensis L. var. barbadensis. Type:
Barbados: locality unknown, [if collected from Description
Barbados, it is extinct there], leg. ign. LINN 1198.1
(lectotype LINN). Ultimate branchlets very slender, 0.81.0mm diam.
Leaves imbricate, appressed or slightly spreading,
Description less markedly in 4 rows; apex obtuse to acute or
mucronate; glands conspicuous.
Ultimate branchlets slender, 1.01.2 mm diam.
Leaves imbricate, slightly spreading, forming 4 reg- Distribution
ular rows of nearly free leaf tips along the branch-
lets; apex obtuse or acute; glands inconspicuous or Bahamas, Cuba (Sierra de Nipe, Cienfuegos, Las
conspicuous. Villas, Isla de la Juventud, Haiti, Jamaica.
TDWG codes: 81 BAH CUB HAI-HA JAM
Distribution
Conservation
Windward Islands: St. Lucia (Petit Piton); extinct on
Barbados. This, the more widespread form of J. barbadensis, is
TDWG codes: 81 WIN-SL nevertheless severely restricted in most of its range
and threatened by habitat changes. Small stands of
Ecology 530 trees are the usual situation in the Bahamas and
in Jamaica (Adams, 1989) and some are also threat-
In its only remaining habitat restricted to limestone ened by cutting as the wood is used in souvenir carv-
cliffs near the summit of a mountain, with other ing. In Cuba, larger populations exist in the Sierra
shrubs. de Nipe and in swampy areas on Isla de la Juventud
[Isla de Pinos], but an accurate assessment of this rounded and entire-hyaline, abaxially bearing 46
taxon in Cuba has not been undertaken to date. flattened, tightly packed pollen sacs. Seed cones
At least some stands in Cuba are in less accessible terminal on erect, short ultimate branchlets, matur-
locations, providing for the time being some mea- ing in one season, becoming irregularly globose or
sure of protection. broad pyriform to nearly reniform, 46 58mm,
IUCN: VU [C2a (i)] usually soft pulpy, resinous, pruinose-blue or dark
purplish blue. Bract-scale complexes 6, decussate,
Uses rarely in 2 whorls, entirely fused, sutures invisible, a
few minute bract tips protruding. Seeds 12(3) per
The wood is locally used for the carving of souvenirs. cone, ovoid-globose, 23mm, more or less keeled,
402 lustrous brown.
Mexico, Durango (Sierra Madre Occidental, summit Juniperus brevifolia (Seub.) Antoine, Cupress.-
area of Cerro Huehuento). Gatt.: 16, t. 2022. 1857. Type: Macaronesia:
TDWG codes: 79 MXE-DU Azores, Flores, [the specimen at P is recorded as
from Faial], C. F. Hochstetter 124 (holotype not
Ecology located, isotypes K, P).
Juniperus blancoi Martnez var. mucronata Spreading or tall shrubs to small trees, to 6 m, dioe-
(R. P. Adams) Farjon, Monogr. Cupressaceae & cious; trunk diam. to 50cm, low branching. Bark
Sciadopitys: 249. 2005. Juniperus mucronata on large trunks with long, soft strips, from reddish
R. P. Adams, Biochem. Syst. Ecol. 28 (1): 158. 2000. brown to grey-brown. Branches numerous, spread-
Type: Mexico: Sonora, Maicoba River, 19 km ing, assurgent or ascending. Foliage dense and stiff,
W of Maicoba, along Mex. Hwy. 16 at km 307, with numerous spreading, jointed ultimate branch-
R. P. Adams 8704 (holotype SRCG). lets, persistent, forming a shrubby, irregular crown
(prostrate at high altitudes). Leaves in alternating
Description whorls of 3, rarely 4, non-decurrent, spreading 4590
degrees at nodes 1.53(5) mm apart, rigid, jointed
Small trees. Leaves on ultimate branchlets rhombic to leaf part adnate with shoot, lanceolate to nar-
or ovoid-rhombic, apiculate; apex just free. Mature rowly ovate (boat-shaped), 410 13mm, usually
roadest near the curved, slightly thickened base;
b of Juniperus brevifolia. Forest remnants are now very
margins entire; abaxial face keeled; leaf colour green restricted and scattered. Afforestation with exotic
to dark green; apex obtuse-acuminate or acute- trees, e.g. Cryptomeria and Eucalyptus, is wide-
pungent; epistomatic, with stomata in two conspicu- spread and causes (irreversible?) habitat changes
ous bands with whitish cuticular wax separated by unsuitable to native trees. Invasive herbs and shrubs,
a midrib, bordered by green margins broader than e.g. Hedychium gardnerianum and Hydrangea mac-
the midrib. Pollen cones solitary, 12 per leaf whorl, rophylla, spread over semi-natural vegetation and
subglobose to ovoid-oblong, 35 24mm; micro- ubiquitous grazing of domestic animals cause fur-
sporophylls 1014, in alternating whorls of 3, pel- ther habitat degradation. Today Juniperus brevifolia
tate, with erose-denticulate margins and acute apex, is only known as a (straggling) shrub, but remains of
bearing 34 abaxial pollen sacs near lower margin. old trunks on some islands indicate its former extent 405
Seed cones maturing in 2 years; mature cones glo- and condition.
bose, 69mm diam., with 1(2) whorls of 3 com- IUCN: VU [A2 (a, c, e); B2ab(ii, iii)]
pletely fused bract-scale complexes, only the upper
whorl fully developed, 3 sutures visible on distal part Uses
of cone, bract tip hidden or very small, tissue soft
pulpy, juicy in most cones; immature cones green, This shrubby juniper is rare in cultivation; it would
hard; mature cones dark red-brown, often slightly be suitable for gardens in areas with mild winters.
glaucous. Seeds 13 per cone, (angular-)ovoid, 56 Its endangered status in the Azores merits wider use
34mm, light brown with obtuse apex. in cultivation especially in connection with ex situ
conservation programmes.
Distribution
USA: W Arizona, California, S Nevada; Mexico: Juniperus cedrus Webb & Berthel., Hist. Nat. Iles
Baja California Norte (including Cedros Island and Canaries 3 (2), Phytogr. Canar. 3 (89): 277. 1847.
Guadalupe Island). Type: Macaronesia: Canary Islands, [Ins. Can.],
TDWG codes: 76 ARI CAL NEV 79 MXI-GU MXN-BC P. B. Webb s.n. (holotype not located, isotype K).
Description Distribution
Trees to 2025 m tall (most surviving plants smaller Canary Islands (Tenerife, La Palma, Gran Canaria,
and shrubby), dioecious; usually monopodial, often Gomera), Madeira.
low branching. Bark on trunks with long, soft strips, TDWG codes: 21 CNY MDR
grey-brown. Branches long, spreading, assurgent or
ascending. Foliage usually sparse, with spreading or Ecology 407
pendulous, jointed ultimate branchlets, persistent,
forming an open, irregular and broad crown (young Originally this species would have been a constitu-
trees more pyramidal). Leaves in alternating whorls ent of the Macaronesian evergreen forest, but as this
of 3, rarely 4, non-decurrent, spreading 3060(90) vegetation type, and particularly the more open,
degrees at nodes 38(10) mm apart, rigid, jointed to drier variant in which Juniperus cedrus had its opti-
leaf part adnate with greenish yellow or glaucous shoot, mum, has been altered or has disappeared altogether,
linear-lanceolate to narrowly ovate (boat-shaped), it is now confined to steep rocky places at altitudes
(4)823 12.3mm, straight, usually broadest near ranging from 450 m to 2400 m a.s.l. Here regenera-
middle; margins entire; abaxial face (weakly) keeled tion can succeed, but a forest with junipers and other
or grooved, green to dark green; apex acuminate or trees will not easily develop.
acute-pungent; epistomatic, stomata in two conspicu-
ous bands separated by a midrib, bordered by green Conservation
margins broader than midrib. Pollen cones solitary,
12 per leaf whorl, subglobose to ovoid-oblong, 36 This species is now restricted to inaccessible places
25mm, orange-brown; microsporophylls 1014, in both in the Canary Islands and Madeira. According
alternating whorls of 3, peltate, with erose-denticulate to David Bramwell, Director of the Botanic Garden
margins, bearing 34 abaxial pollen sacs near lower in Las Palmas, there are probably not more than 100
margin. Seed cones axillary on short (12mm) dwarf mature trees remaining in the wild (on Gran Canaria
shoots with whorls of minute scale leaves, maturing in less than 6, numbers on Las Palmas are unknown).
2 years; mature cones globose, 814mm diam., with Grazing by goats prevents successful regeneration
1(2) whorls of 3 completely fused bract-scale com- and it is hardly possible to speak of viable popula-
plexes and only the upper whorl fully developed, 3 tions in most places.
sutures visible on distal part of cone; bract tip hidden IUCN: EN [B2ab (ii, iii, v); C2a (i)]
or very small; tissue soft pulpy, more or less resinous;
immature cones green, hard, mature cones orange- Uses
brown. Seeds 13 per cone, ovoid, 510 46mm.
When Canary Island juniper was still abundant its
Taxonomic notes wood was highly sought after due to its durability;
it was used for fence posts and for furniture mak-
The variation found in the size and shape of the ing. This species is rare in cultivation and ought to
leaves of this species is considerable, from narrow be tried more (perhaps not in countries with rela-
(ca. 1mm) and long (23mm), pungent to broad tively few hot, sunny days in summer). It grows on
(2.2mm) and short (8mm), obtuse. As in J. oxyce- Tenerife at altitudes where frost and snow are not
drus, to which it is somewhat more distantly related, uncommon, but solar radiation is intense. Ex situ
length and acuteness of the leaves generally decrease conservation is urgently needed to build up stock
with increasing age of the trees and probably with for future replanting and horticulture could have an
increasing exposure to light and weather. Specimens important role to play in this effort.
Juniperus chinensis L., Mant. Pl. 1: 127. 1767. pulpy, resinous, light brown, red-brown or glaucous
cones. Bract-scale complexes (4)6(8), decussate,
Etymology occasionally ternate, usually the lower pair infertile,
entirely fused, sutures of upper scales partly visible
The species epithet means from China. as a curved ridge terminating in a minute bract tip.
Seeds (1)24(5) per cone, ovoid-flattened, 37
Vernacular names 26mm, with small resin pits and longitudinal,
shallow grooves, light yellowish brown.
Chinese juniper; yuan bai (Chinese); ibuki,
byakushin, ibuki-byakushin, kamakura-byakushin Taxonomic notes
408 (Japanese); Kong Nam Tsong (Korean)
Juniperus chinensis is one of the most variable spe-
Description cies in the genus. Its polymorphy has been the cause
of an almost inextricable synonymy, especially so
Decumbent, sometimes ascending or erect shrubs, because horticulturists have described taxa at vari-
or a tall trees to 2025 m with a monopodial, erect ous ranks from variations observed in cultivation
trunk to 1 m diam., dioecious or rarely monoecious. which should have been treated as cultivars.
Bark on tree trunks fissured, exfoliating in long,
fibrous, thin strips, becoming darker grey-brown. Distribution
Branches spreading horizontally or ascending, in
shrubs spreading, ascending or erect, spreading Myanmar [Burma], China, Japan, Korea, Russian
or subpendulous in trees, forming dense matting Far East, Taiwan.
crowns in decumbent shrubs to initially pyramidal, TDWG codes: 31 KHA KUR PRM SAK 36 CHC-CQ
then open and irregular crowns in trees. Foliage CHC-GZ CHC-HU CHC-SC CHC-YN CHI-NM
branchlets numerous, spreading, assurgent or erect CHI-NX CHM-HJ CHM-JL CHM-LN CHN-BJ
in shrubs, spreading to pendulous in trees, slen- CHN-GS CHN-HB CHN-SA CHN-SD CHN-SX
der, 11.5mm diam., subterete to quadrangular in CHN-TJ CHS-AH CHS-FJ CHS-GD CHS-GX CHS-HE
branchlets with scale leaves, persistent. Leaves decus- CHS-HK CHS-HN CHS-JS CHS-JX CHS-ZJ CHT 38
sate or in alternating whorls of 3, short decurrent, JAP-HK JAP-HN JAP-KY JAP-SH KOR-NK KOR-SK
of two types: needle-like and scale-like. Needle-like TAI 41 MYA
leaves mostly ternate, linear-subulate,(4)610(12)
0.50.7mm, pungent, adaxially canaliculate, epi- Ecology
stomatic, with two narrow bands of stomata. Scale
leaves increasingly dominant with age of plant in In a few localities this widespread species forms
most varieties, always decussate, imbricate, closely groves of tall trees (e.g. in S Gansu), or it is mixed
appressed, on ultimate branchlets 1.53 1mm, with pines and deciduous angiosperms at canopy
ovate-rhombic to rhombic-lanceolate, abaxially level. It is much more common, under conditions
with a conspicuous central, elliptic gland, green largely determined by mans agricultural practices,
or glaucous green; margins entire; apex obtuse to in secondary vegetation, on open, rocky slopes. The
acute; stomata few at base abaxially, in two concave altitudinal range is (100)14002400(2700) m a.s.l.
fields separated by a narrow midrib adaxially. Pollen Widespread planting and subsequent establishment
cones terminal on ultimate branchlets with scale- in areas where it was not originally native have made
like or acicular leaves, ovoid-oblong, 46 23mm; it difficult to establish its original habitat and types
microsporophylls 1218, decussate, peltate-cordate, of vegetation. While now predominantly montane,
convex, upper margin entire, abaxially bearing 34 it may have been part of lowland forests in the past
ovoid-oblong pollen sacs. Seed cones terminal on (Wang, 1961). High montane varieties J. chinensis
erect, short shoots 310mm long with small scale var. sargentii and var. tsukusiensis occur on rocky
leaves, maturing in second season to (irregularly) outcrops and amongst boulders and have attained a
globose or broad pyriform, 410mm, usually dry decumbent habit.
Uses seed cones only on ultimate branchlets with scale
leaves.
Juniperus chinensis is (with Platycladus orientalis)
one of the two most commonly planted cupressa- Distribution
ceous trees in traditional Chinese gardens, such as
around temples and in the extensive grounds of the Myanmar [Burma]; China; Japan; Korea; Taiwan.
Forbidden City in Beijing. These grounds are now TDWG codes: 36 CHC-CQ CHC-GZ CHC-HU
virtual reserves for large specimen trees, which have CHC-SC CHC-YN CHI-NM CHN-BJ CHN-GS CHN-HB
all but disappeared from the countryside. Its wood is CHN-SA CHN-SD CHN-SX CHN-TJ CHS-AH CHS-FJ
highly valued for furniture making and joinery and CHS-GD CHS-GX CHS-HE CHS-HK CHS-HN CHS-JS
is hard and durable. In horticulture, J. chinensis has CHS-JX CHS-ZJ 38 JAP-HK JAP-HN JAP-KY JAP-SH 409
been the source of many cultivars and (as J. sphaerica KOR-NK KOR-SK TAI 41 MYA
Lindl.) one parent of a purported hybrid (of garden
origin) with J. sabina, for which Van Melle (1946) Conservation
proposed the name J. media, a later homonym of
J. media V. D. Dmitriev (1938) = J. semiglobosa. A IUCN: LC
proposal to conserve van Melles name failed and the
cultivars thought to derive from this hybrid origin
are now to be listed under J. pfitzeriana based on Juniperus chinensis L. var. sargentii A. Henry, in
a selection made by the Spth Nursery in Berlin, Elwes & Henry, Trees Gr. Brit. Ireland 6: 1432.
Germany in the 1890s (Pfitzer Junipers). A study 1912. Juniperus sargentii (A. Henry) Takeda ex
using RAPDs (Le Duc, Adams & Zhong, 1999) sup- Nakai, Bot. Mag. (Tokyo) 44: 511. 1930. Type:
ports the notion of a hybrid origin with parents Japan: Hokkaido, Kitami Prov., Rebun-jima,
J. chinensis and J. sabina of the Pfitzer cultivars, M. Furuse 9385 (neotype K). Fig. 134
for which the correct botanical name is Juniperus
pfitzeriana (Spth) Schmidt [Pfitzer Group]. This Vernacular names
species is also of major importance in bonsai and
penjing culture. yan bai (Chinese); Miyama-byakushin (Japanese)
Description Etymology
Decumbent or low shrubs to 80100cm, often pros- The species epithet refers to the Mexican State of
trate or creeping over rocks. Foliage very dense, ulti- Coahuila.
mate branchlets 825mm long, slender, ca. 1mm
thick, quadrangular in cross-section, lax but not Vernacular names
drooping. Leaves on most shrubs of two types, nee-
dle-like and scale-like, or only scale leaves; acicular No common names are recorded for this species.
leaves in whorls of 3, 48mm long; scale leaves 11.3
1mm, triangular rhombic or gibbous, obtuse or Description
slightly apiculate. Gland central, small, ovoid, con-
spicuous but inactive. Seed cones small 45mm Large shrub to 5 m, or small tree to 10 m, dioe-
diam., red-brown to purplish brown, with some cious, rarely monoecious; multistemmed or with
glaucous bloom. a short trunk and a maximal diam. of 50cm. Bark
of thick branches and trunk fibrous, exfoliating
Distribution in strips, weathering ash-grey. Branches numer-
ous, long, ascending from base or spreading, often
China?, Taiwan (Chingshui Cliffs), S Japan curved, forming a broad, rounded or more irregu-
(Yakushima). lar and open crown. Foliage branches numerous,
TDWG codes: 38 JAP-KY TAI irregularly disposed but not pendulous, ultimate
ranchlets spreading 3070 degrees, slender and
b and, of course, it is found in rocky areas adjacent to
stiff, up to 15mm long, 11.5mm thick, (weakly) grasslands, often in association with Opuntia spp. or
quadrangular in cross section, covered with scale Yucca spp. It is unusual in that J. coahuilensis sprouts
leaves, persistent. Leaves decussate on all ultimate from the stump when cut or burned. This feature has
branchlets, decussate or in alternate whorls of 3 on probably allowed it to remain in the grasslands in
older branchlets, imbricate, scale-like, 1.21.7(1.8) spite of periodic grass fires that kill all other juni-
0.71.2mm on lateral branchlets, decurrent, rhom- per species [R. P. Adams, pers. comm.]. It is consid-
bic or ovoid-rhombic; apex appressed or rarely ered to be weedy in some areas by sheepmen and
free, obtuse to acute; margins minutely denticulate; cattlemen.
stomata on abaxial side limited to two small spots
near leaf base, on adaxial surface in two tapering Conservation 411
bands; glands central on small scale leaves, more
or less conspicuous, large, rounded to oblong, flat IUCN: LC
or slightly raised, exudate often present; leaf colour
yellowish green or light green. Pollen cones numer-
ous, terminal, solitary, ellipsoid or ovoid-oblong, Juniperus comitana Martnez, Anales Inst. Biol.
34 22.5mm; microsporophylls 810, decussate, Univ. Nac. Mxico 15 (1): 12. 1944. Type: Mexico:
peltate, acute, with minutely denticulate upper mar- Chiapas, Comitn de Domnguez, 12 km S of
gins and bearing 34 abaxial pollen sacs. Seed cones Comitn, M. Martnez 6700 (holotype MEXU).
terminal, maturing in one season attaining variable
colours from pinkish(red) or orange(red) to pur- Etymology
plish blue, usually glaucous, (sub-)globose or some-
times broad ovoid, 57mm diam. (if ovoid to 9mm The species epithet refers to the town of Comitn in
long), internally soft pulpy, succulent, yellowish to Chiapas, Mexico.
orange-brown. Bract-scale complexes 4(6), entirely
fused, decussate; bract apices minutely exserted or Vernacular names
hidden; surface smooth, rugose when dry. Seeds
1(2) per cone, ovoid-globose, 35(6) 34(5) No common names have been recorded for this
mm, with rounded base and acutish apex, shallowly species.
grooved and ridged on sides, not or slightly resinous,
chestnut brown, with a large, lighter lobed hilum Description
proximally.
Trees to 810 m, presumably dioecious; trunk mono-
Distribution podial, erect or forked and twisted, usually branch-
ing well above base, up to 6080cm d.b.h. Bark on
Mexico: Chihuahua, Coahuila, Durango, Nayarit, trunk 510mm thick, fibrous, lacerated, exfoliating
Tamaulipas, Zacatecas; USA: SW Texas. in long strips exposing purplish inner bark; outer
TDWG codes: 77 TEX 79 MXE-CO MXE-CU bark red-brown to grey-brown. Branches long,
MXE-DU MXE-TA MXE-ZA MXS-NA spreading or ascending, foliage forming in mature
trees a broad, rounded or irregular crown. Foliage
Ecology branches numerous, forming lax, dense foliage tufts,
irregularly disposed, ultimate branchlets of irregular
This species occurs in the high desert (12002000 m length from 420(25) mm, very slender, 0.71mm
a.s.l.) Bouteloua grasslands and adjacent rocky areas. diam., more or less quadrangular in cross-section
It is unusual, for Juniperus, in that it occurs in these to more terete on older branchlets which become
seemingly undisturbed grasslands. In Mexico, the rough with free leaf apices, persistent. Leaves on
populations may occur in canyons or on alluvial fans ultimate and lower lateral branchlets decussate,
but occasionally on slower growing sections of Guatemalan Sierra de los Cuchumatnes where both
older branchlets in alternating whorls of 3, closely J. comitana and J. standley still occur.
appressed, on ultimate branchlets ovoid-rhombic or IUCN: EN [B2ab (ii, iii, v)]
rhombic-triangular, 11.4 0.71mm; margins hya-
line and minutely erose to serrulate; apex acuminate Uses
or sometimes obtuse; stomata few abaxially near
base along margins, abundant adaxially from base The wood of this juniper is used locally by the
to apex; glands small, usually inactive; leaf colour Amerindian population for firewood and also for
light green. Pollen cones terminal, solitary, oblong, fence posts.
46mm long; microsporophylls 1012, decussate,
412 peltate, with minutely hyaline-denticulate upper
margins, bearing (3)4 abaxial pollen sacs. Seed Juniperus communis L., Sp. Pl. 2: 1040. 1753.
cones numerous, terminal, solitary on very short
lateral branchlets, maturing in one year to purplish Etymology
brown cones with a glaucous bloom; mature cones
subglobose, 57mm diam. Bract-scale complexes 4, The species epithet means common; Linnaeus
decussate, lower pair smaller than upper pair meet- knew it in abundance from Sweden.
ing at cone apex, completely fused, with smooth
external surface, lustrous when ripe; 2 decussate Vernacular names
pairs of bract tips visible, but these less than 1mm
long, in ripe cones sometimes submerged; internal Common juniper; Genvrier commun (French);
tissue soft pulpy, resinous. Seeds usually 1 per cone, Gemeine Wacholder (German); Mozhzhevelnik
ovoid, 45(6) 3.54mm. obyknovennyy (Russian)
Distribution Description
Europe, North Africa, Caucasus, Siberia to Russian Dwarf juniper, Prostrate juniper
Far East, Western Asia, Central Asia.
TDWG codes: 10 DEN FIN FOR GRB ICE IRE Description
NOR SWE 11 AUT-AU AUT-LI BGM-BE BGM-LU
CZE-CZ CZE-SK GER HUN NET POL SWI 12 BAL Decumbent or spreading shrubs to 1.5 m, rarely
COR FRA-CI FRA-FR POR SAR SPA-AN SPA-SP a large shrub or small tree to 10 m, not forming a
13 ALB BUL GRC ITA-IT ITA-SM KRI ROM SIC-SI conical or pyramidal crown; branchlets ascending at
TUE YUG-BH YUG-CR YUG-KO YUG-MA YUG-MN tips. Leaves at nodes 36(10) mm apart, spreading
YUG-SE YUG-SL 14 BLR BLT-ES BLT-KA BLT-LA at angles of 3070 degrees, often ascending or more
BLT-LI KRY RUC RUE RUN RUS RUW UKR-MO or less erect, at internodes of c. 5mm, curved at base
UKR-UK 20 ALG 30 ALT BRY CTA IRK KRA TVA (sometimes straight), linear, (5)1020 11.7mm;
WSB YAK 31 AMU PRM SAK 33 NCS TCS 34 AFG CYP stomatal band from as wide as each green margin to
EAI IRN IRQ LBS-LB LBS-SY TUR 37 MON nearly twice as wide. Seed cones 69mm, globose.
Seeds 3 per cone.
Ecology
Distribution
This, the typical variety, is largely a pioneer wood-
land species, occupying natural rock outcrops and North America, from Alaska to Newfoundland, S to
other places with skeletal soil and abundant sunlight Arizona and South Carolina.
in woodland and light forest, both broad-leaf and TDWG codes: 70 ASK NUN NWT YUK 71 ABT BRC
coniferous forest (especially Pinus sylvestris-Betula MAN SAS 72 LAB NBR NFL-NE NFL-SP NSC ONT
spp.-Quercus spp.), in which it can obtain local PEI QUE 73 COL IDA MNT ORE WAS WYO 74 ILL
dominance after disturbances (non-fire). It is also IOW KAN MIN MSO NDA NEB OKL SDA WIS 75 CNT
prevalent in the ecotone between open woodland INI MAI MAS MIC NWH NWJ NWY OHI PEN RHO
and grassland on poor sandy soils and on stabilised VER WVA 76 ARI CAL NEV UTA 77 NWM 78 GEO
inland sand dunes. It occurs often with Calluna vul- KTY NCA SCA TEN VRG
Ecology barrens, commonly in the ecotone between open
vegetation and coniferous forest with Abies balsamea
In a wide range of habitats from sea shores to (sub) or Picea rubens.
alpine meadows and rocky ridges. Mostly associated
with coniferous woodland and forest, but usually Conservation
occupying localities or topography less suitable for
tree growth, or in open woodland on thin or poor IUCN: LC
sandy soil or dolomite. Associated e.g. with Pinus
banksiana, P. ponderosa, P. flexilis, Abies balsamea, A.
lasiocarpa, Picea engelmannii, P. glauca, and Populus Juniperus communis L. var. nipponica (Maxim.)
tremuloides in lowland to subalpine forests, with E. H. Wilson, [Conifers & Taxads Japan] Publ. 415
shrubs and herbs in coastal or inland sand dunes, in Arnold Arbor. 8: 81. 1916. Juniperus nipponica
(sub)alpine meadows, or in peat bogs. The altitudinal Maxim., Bull. Acad. Imp. Sci. Saint-Ptersbourg 12:
range is from 5 m to 3800 m a.s.l. It is indifferent to 230. 1868; Juniperus rigida Siebold & Zucc. subsp.
soil types, with pH from very acid to neutral or mildly nipponica (Maxim.) Franco, Bol. Soc. Brot., ser. 2,
alkaline and with low or fluctuating water tables. 36: 119. 1962. Type: Japan: Honshu, Iwate Pref.,
[Prov. Nambu, in alpibus altioribus], Tschonoski
Conservation (Chnosuka Sugawa) [ex herb. C. J. Maximowicz
s.n.] (holotype LE).
IUCN: LC
Vernacular names
plate 15. Juniperus drupacea. 1. Habit of tree. 2. Branch with foliage. 3. Leaves. 4. Cross-section of leaf.
5. Pollen cone. 6, 7. Seed cones (with cross-section showing fused seeds).
subgenus Caryocedrus for it. The partly fused seeds semi-natural. They suggested protective status for at
stressed by Endlicher as a diagnostic character of least one of the major stands in Greece.
this section (= J. drupacea), as well as the cone itself, IUCN: LC
require a careful ontogenetic study for which unfor-
tunately no material was available. What is clear is Uses
that the ovules originate independently in the axils
of the fertile whorl of 3 bracts, so that fusion is a sec- The wood of large trees is valuable as timber with
ondary character state. Some similarities with large decay resistant properties but it is not extensively
cones of J. oxycedrus subsp. macrocarpa (in which used for this purpose. In Turkey, the local population
the 3 seeds remain free but lie closely appressed so harvest the fleshy cones, which have a high content
as to determine each others shape during growth) of sugars, for consumption as a kind of marmelade 423
suggest that these differences are relative rather than or dried fruit. Its habit in cultivation tends to remain
absolute. columnar for many years and its large leaves with
conspicuous stomatal bands are also ornamental.
Distribution Yet its use in gardens is limited despite good growth
rates and tolerance to frost; this omission deserves
Greece, Turkey (mountains along Mediterranean to be rectified as it is probably a better species for
coast), Syria, Lebanon, Israel. horticulture than the more common J. oxycedrus
TDWG codes: 13 GRC 34 LBS-LB LBS-SY PAL-IS (in southern Europe and regions with equivalent
TUR climate).
Ecology
Juniperus durangensis Martnez, Anales Inst. Biol.
Juniperus drupacea occurs in mixed (low) mon- Univ. Nac. Mxico 17 (1): 94. 1946. Type: Mexico:
tane conifer forest and woodland with Pinus bru- Durango, El Salto, Puerto de Santo Domingo, M.
tia, P. nigra, Abies cephalonica, A. cilicica, Cedrus Martnez 7015 (holotype MEXU).
libani, Juniperus excelsa, J. foetidissima, J. oxycedrus,
Quercus coccifera, Q. ilex, and more rarely with Fagus Etymology
orientalis. Forest degradation has changed the vege-
tation in some areas to maquis in which the junipers The species epithet refers to the State of Durango in
have regenerated or spread. It occupies, with other Mexico.
junipers, the rockier, more shallow soils, often on
calcareous but also on granitic rock. The altitudinal Vernacular names
range is from 600 m to 1800 m a.s.l. The climate is
Mediterranean, with winter rain, sometimes snow, Durango juniper; tscate (Spanish)
and dry, hot summers.
Description
Conservation
Shrubs or small trees to 6 m, presumably dioe-
This species is rare in Greece, Lebanon and Israel, cious; trunk forked at base or branching low, or
where it is probably Vulnerable (Greece) or even erect and curved or contorted, to 3040cm diam.
Endangered (Israel); however in Turkey and Syria Bark becoming fibrous, 510mm thick, exfoliating
it is still more widespread and locally common. in long, thin strips, light brown weathering grey.
Boratynski & Browicz (1983) gave an account of the Branches thick, contorted, spreading and assurgent,
populations in Greece, all in the Parnon Massif in foliage dense, forming an irregular and open crown
the Peloponnisos. While some stands represent with wide spreading branches. Foliage branches
degraded fir forest (Abies cephalonica) or have been numerous, rigid, forming dense tufts, ultimate
converted to pasture, others appear to be natural or branchlets short, rigid, spreading in all directions,
straight or towards ends of foliage branches down- herbarium collections examined, a habitat consisting
curved, 410(18) mm long, 1.21.5mm diam., more of (cleared) oak-juniper woodland was mentioned,
or less quadrangular in cross-section, leading ulti- with Juniperus deppeana var. robusta also present.
mate branchlets to 2mm diam. Leaves decussate on According to Adams (2011) it is found on rhyolite
ultimate lateral branchlets, in alternating whorls of 3 rocks and in openings of pine-oak forests, and his
on lower branchlets, imbricate, on ultimate branch- book shows similar habitat photographs. This habitat
lets appressed, broadly ovoid-rhombic, 0.81.2 is confirmed on the labels of several herbarium col-
0.81mm, often nearly as wide as long, gibbous; lections kept in ARIZ (Tucson, Arizona). These col-
margins thick, minutely denticulate; apex obtuse, lections were made at altitudes of 15002700 m a.s.l.
leaves on older branchlets broad rhombic, acute;
424 stomata few on lower margin abaxially and scattered Conservation
stomata from base to apex adaxially; gland central,
small, oval-oblong to linear, inactive; leaf colour Although it is known from a wide range in N Mexico,
dark green, new foliage often yellowish green. Pollen only a limited number of verified collections have
cones terminal, solitary, subglobose or ovoid, 2.53 been made and it is apparently not common. Often,
2mm; microsporophylls 1012, decussate, peltate, these collections were made in clearings of pine-
as broad as long, with hyaline-denticulate margins, oak forest or woodland, some of these appear to be
obtuse, bearing 23 abaxial, relatively large pollen natural rocky areas with few trees. A more detailed
sacs. Seed cones numerous, terminal on curved or survey of specimens has now established its approxi-
straight, 23mm long branchlets; mature cones sub- mate extent of occurrence and probable conserva-
globose to nearly reniform, often wider than long, tion status. It does not seem to be threatened at
46 57mm, irregular, more or less bilobed if with present.
2 seeds, smooth (rugose in sicco), orange-brown or IUCN: LC
reddish brown, often glaucous to pruinose. Bract-
scale complexes 4, decussate, upper pair larger than Uses
lower pair, bract tips exserted, 0.5mm, triangular
and more or less curved; tissue soft pulpy, succulent No specific uses have been recorded of this species;
or sometimes dry. Seeds 13(4) per cone, variable it presumably serves both as firewood and for fence-
but mostly angular-ovoid, 34.5 23mm, curved posts locally.
or not, ridged and/or shallowly pitted, hilum a third
to half of seed, resinous or not, seed coat light brown
or reddish brown, but hilum lighter and usually Juniperus excelsa M.-Bieb., Tabl. Prov. Mer.
turning yellowish brown. Casp.: 204. 1798. [Beschr. Lnd. Terek & Kur, Bot.
Anhang: 204. 1800.]
Distribution
Etymology
Mexico: Aguascalientes, Chihuahua, Durango,
Jalisco, Sonora, Zacatecas. The species epithet means high and perhaps refers
TDWG codes: 79 MXE-AG MXE-CU MXE-DU to the considerable size of this species compared to
MXE-ZA MXN-SO MXS-JA the shrubby species known from central and west-
ern Europe.
Ecology
Vernacular names
Martnez (op. cit.) described the habitat of this spe-
cies as very poor, dry, rocky soil, and presented some Greek juniper, Grecian juniper, Crimean juniper,
photographs of trees that are apparently growing in Turkestan juniper (subsp. polycarpos); Ard, Boylu
a deforested landscape, which probably had been ard (Turkish); Dedali-gwia (Iran); Parmiro ala
covered with pine-oak woodland. On two labels of (Central Asia, subsp. polycarpos)
Description TCS-GR 34 AFG CYP IRN LBS-LB IRQ LBS-SY TUR
35 OMA 40 PAK WHM-JK
Trees (rarely shrubs) to 2025 m, monoecious or more
commonly dioecious; monopodial; trunk up to 2.5 m Ecology
d.b.h., or a (decumbent) multistemmed shrub. Bark
on trunks exfoliating in longitudinal fibrous strips, In shrubland (Mediterranean), deciduous wood-
reddish brown weathering grey-brown. Branches land and coniferous forest, in the interior eastern
spreading or ascending, foliage branches very dense, parts of its range also in montane Seriphidium mari-
short, stiff and spreading, or longer and more or less timum (Artemisia maritima) steppes and on alpine
lax, forming a dense, rounded or irregular, sometimes scree or moraines. The altitudinal range is (100
sympodial crown. Foliage branchlets ultimately sub- )5003000(3950) m a.s.l. It is most common on 425
terete or weakly quadrangular, 0.71.3mm wide, cov- dry slopes of calcareous hills and mountains in the
ered with closely appressed leaves, persistent. Leaves Mediterranean climate parts of its range, but occurs
on mature plants normally scale-like, decussate, also on siliceous rock in the continental interior,
imbricate, decurrent, 0.61.6 0.40.9mm, ovate- more often near intermittent groundwater sources.
rhombic, obtuse; margins entire; stomata in 2 incon- In Central Asia, it is restricted to the more western
spicuous lines on each side mostly near base; leaves parts of the numerous mountain ranges, where it
abaxially glandular; gland central, large and conspic- forms, with some other species, (J. semiglobosa, J.
uous, elliptic or nearly circular, often with exudate; pseudosabina, J. sabina) extensive juniper forest or
leaf colour yellowish green to dull light green. Pollen woodland (Archa forest) on all but the N-facing
cones numerous, solitary, terminal on short branch- slopes (where e.g. Picea schrenkiana dominates, a
lets, globose to subglobose, 34 23mm; microspo- conifer that is more abundant and not restricted to
rophylls 810, decussate, peltate, with rounded entire N-facing slopes farther east). In contrast to North
margins, bearing 34 abaxial pollen sacs. Seed cones America, this juniper forest is not mixed with pines
terminal on short, erect branchlets, maturing in the (Pinus spp.) and angiosperms are uncommon in it.
second season and becoming (sub)globose, 612( Several species of Juniperus, including low shrubs,
14) mm diam., pinkish brown, blue or purplish blue, therefore determine the structure of these forests or
usually glaucous or pruinose. Bract-scale complexes woodlands in large areas, often from the valley bot-
46(8), decussate, entirely fused, sometimes sutures tom to the tree line.
faintly visible near apex of cone, bract tip small, ca.
0.5mm, on a small ridge, scale tissue dry pulpy or Uses
rarely more succulent, resinous. Seeds (2)36(8)
per cone, subovoid-conical, more or less flattened or The wood of J. excelsa is durable and hard and, in
curved, 46 34mm, yellowish- to reddish brown, parts of Turkey at least, sufficiently common to be
with lighter hilum at base. utilized. However, trees grow slowly and take two
centuries or more to attain any harvestable size, so
Distribution its exploitation for timber is or would not be sustain-
able if it occurred on a commercial scale. In many
SE Europe: Albania, Bulgaria, Greece, Macedonia; parts of SW Asia it also serves as firewood, especially
E Europe: Ukraine: Krym [Crimea]; Caucasus: for people who live (temporarily) in the mountains.
Armenia, Azerbaijan, Gruzija, Severo-Osetiya, The wood of large trees is used for carpentry and
Russia(?); Central Asia: Kazakhstan, Kirgyzstan, furniture. Foliage is sold and burnt as incense. Greek
Tadzhikistan, Turkmenistan, Uzbekistan; W Asia: juniper is uncommon in horticulture and only a few
Afghanistan, Cyprus, Iran, Lebanon, Syria, Turkey; cultivars have been selected; the one best known in
Arabian Peninsula: Oman; S. Asia: NW India, Britain is a columnar form with glaucous and more
Pakistan. juvenile type foliage properly named Stricta but
TDWG codes: 13 ALB BUL GRC YUG-MA 14 KRY 32 also known under several varietal names. Forms
KAZ KGZ TKM TZK UZB 33 NCS-SO TCS-AR TCS-AZ with pendulous foliage and with dwarf growth have
also been named and are used more often in central Juniperus excelsa M.-Bieb. subsp. polycarpos
and southern Europe. (K. Koch) Takht., Fl. Yerev.: 53. 1972. Juniperus
polycarpos K. Koch, Linnaea 22: 303. 1849;
2 subspecies are recognized: Juniperus excelsa M.-Bieb. var. polycarpos (K. Koch)
Silba, Phytologia Mem. 7: 34. 1984. Type: Turkey:
Gmshane, Anadolu Daglari, Taltaban, P. E. E.
Juniperus excelsa M.-Bieb. subsp. excelsa. Type: Sintenis 5520 (neotype L). Pl. 16
Ukraine: Crimea, Krymskiye Gory, P. S. Pallas,
[ex herb. Pallas] s.n. (lectotype LE). Juniperus macropoda Boiss., Fl. Orient. 5: 709. 1884.
Juniperus seravschanica Kom., Bot. Zurn. (Moscow
426 Description & Leningrad) 17: 481. 1932; Juniperus polycarpos
K. Koch var. seravschanica (Kom.) Kitam., [Fl. Pl.
Trees to 2025 m; ultimate branchlets slender, W. Pakist. Afghan.: 7. 1964] Add. & Corr. Fl. Afghan.:
0.71mm diam., weakly quadrangular or subterete, 68. 1966; Juniperus excelsa M.-Bieb. subsp. seravs-
often in regularly disposed sprays, more or less lax. chanica (Kom.) Imkhan., Bot. Zurn. 75 (3): 407. 1990.
Seed cones 614mm diam., mostly darker hues, Juniperus turcomanica B. Fedtsch., in Fedtschenko
glaucous or pruinose. Seeds 36(8) per cone. et al., Fl. Turkmenii 1: 14. 1932; Juniperus excelsa
M.-Bieb. subsp. turcomanica (B. Fedtsch.) Imkhan.,
Distribution Bot. Zurn. 75 (3): 408. 1990; Juniperus polycarpos
K. Koch var. turcomanica (B. Fedtsch.) R. P. Adams,
SE Europe: Albania, Bulgaria, Greece, Macedonia; Phytologia 86 (2): 50. 2004.
E Europe: Ukraine: Krym [Crimea]; Caucasus: Juniperus polycarpos K. Koch var. pendula Mulk.,
Armenia, Azerbaijan, Gruzija, Severo-Osetiya; Dokl. A. N. Armen. S.S.R. 45 (2): 86. 1967; Juniperus
W Asia: Cyprus, Iran (Elburz Mts.), Iraq, Lebanon, excelsa M.-Bieb. subsp. polycarpos (K. Koch) Takht.
Syria, Turkey; Central Asia: Turkmenistan (Kopet var. pendula (Mulk.) Imkhan., Bot. Zurn. 75 (3): 407.
Mts.). 1990.
TDWG codes: 13 ALB BUL GRC YUG-MA 14 KRY 32
TKM 33 NCS-SO TCS-AR TCS-AZ TCS-GR 34 CYP Description
IRN IRQ LBS-LB LBS-SY TUR
Trees to 20 m, rarely (decumbent) shrubs; ultimate
Ecology branchlets thicker (11.3mm diam.), subterete, rigid,
spreading in all directions. Seed cones 812(14)
This subspecies occurs in shrubland (Mediterranean), mm diam., pinkish brown to purplish blue, usually
woodland and pine forest on dry hillsides and rocky pruinose. Seeds (2)34(6) per cone.
mountain slopes at altitudes from 100 m to 2000(
2700) m a.s.l. It is associated with Pinus brutia, Distribution
P. nigra, Cedrus libani, Abies cephalonica, A. cilicica,
A. nordmanniana, Juniperus foetidissima, J. oxyce- Central Asia: Kazakhstan, Kirgyzstan, Tadjikistan,
drus, Quercus spp., Pistacio lentiscus, Berberis, Acer Turkmenistan, Uzbekistan; Caucasus: Armenia,
obtusifolium, etc. It grows most often on limestone Azerbaijan, Russia (?); W Asia: Afghanistan, Iran, E
and also on igneous rocks, but not in acidic soil. The Turkey; Arabian Peninsula: Oman; S Asia: Pakistan,
climate is Mediterranean, with winter rain and dry, NW India (Kashmir).
hot summers, grading eastward into more conti- TDWG codes: 32 KAZ KGZ TKM TZK UZB 33
nental conditions, where J. excelsa subsp. polycarpos TCS-AR TCS-AZ TCS-GR 34 AFG IRN TUR 35 OMA
predominates. 40 PAK WHM-JK
Conservation
IUCN: LC
427
plate 16. Juniperus excelsa subsp. polycarpos. 1. Habit of tree. 2. Branch with foliage and seed cones.
3. Branchlet with adult leaves. 4. Branchlet with juvenile leaves. 5. Seedling. 6, 7. Pollen cones. 8. Branchlet
with seed cones. 9. Seed cone. 10. Opened seed cone with seeds.
Ecology ing a sympodial, broad, rounded crown. Foliage
branchlets ultimately slender, flaccid, subterete,
The continental subspecies occupies a range of 11.2mm wide, covered with imbricate leaves, per-
habitats from montane (mixed) coniferous forest to sistent. Leaves on mature plants scale-like, decus-
upper montane steppes dominated by Seriphidium sate, imbricate, decurrent with free to spreading
maritimum (Artemisa maritima). Its altitudinal apex, 24 0.71mm, broad lanceolate to rhombic,
range is (550)10003300(3950) m a.s.l. In conifer- acute to acuminate-pungent; gland abaxially in cen-
ous forest it can be associated with Pinus gerardiana, tral depression, elliptic or oblong; margins nearly
P. wallichiana, Cedrus deodara or Abies pindrow, entire-hyaline or minutely denticulate; leaf colour
in broad-leaved forest or woodland with Carpinus yellowish green to light green; stomata abaxially in
428 sp., Juglans nigra, Malus sieversii, Crataegus sp., 2 inconspicuous lines near the base, adaxially in two
Prunus sp., Sorbus sp. and shrubs e.g. Dodonaea and bands barely separated by a midrib. Pollen cones
Sageretia. At its highest altitudes in Afghanistan and numerous, solitary, terminal on short branchlets,
Pakistan it forms very open stands on bare scree or ovoid-oblong, 35 1.52mm, often elongated at
glacial moraines with decumbent Juniperus squa- pollen dispersal; microsporophylls 1016, decus-
mata. It is found on calcareous as well as siliceous sate, peltate-acuminate, with hyaline-erose margins,
rocks, usually in shallow, stony soils, but sometimes bearing 24 abaxial pollen sacs. Seed cones terminal
in loess or loam, and on dry slopes or near intermit- on very short, erect lateral branchlets, maturing in
tent streams. the second season to (sub)globose, 8.516 (in one
var. up to 20, in another down to 6) mm diam., light
Conservation brown, usually glaucous or pruinose cones. Bract-
scale complexes 6, decussate, entirely fused, sutures
IUCN: LC often visible on entire cone forming a polygonal pat-
tern; bract tip usually conspicuous, triangular, ca.
1mm, on a small transverse ridge; scale tissue dry
Juniperus flaccida Schltdl., Linnaea 12: 495. 1838. pulpy. Seeds (4)68(12) per cone, angular-conical,
more or less flattened, 34mm long, reddish brown,
Etymology with lighter hilum at base.
Uses Description
No uses have been recorded for this species; the Leaves with hyaline, (minutely) denticulate margins,
wood may locally serve for fuel or be used for fence acuminate-pungent, glaucous green. Seed cones 429
posts. small, 68mm diam., with only 13 seeds.
Description Conservation
Foliage spreading to pendulous, not in more or Only known from a limited number of locations in
less flattened, distichously branching sprays, green. which forest cover is decreasing overall, this variety
Leaves with nearly entire-hyaline margins, acumi- could be at some risk of extinction.
nate-pungent. Seed cones 8.515mm diam., with IUCN: VU [B2ab (ii, iii, v)]
more than 3 and up to 12 (usually 68) seeds.
This species was named after Rafael P. Gamboa, at This species was recently found to be closely related
the time Governor of the State of Chiapas, Mexico. to Jupiperus deppeana in an analysis of DNA
sequence data (Adams & Schwarzbach, 2006) and
Vernacular names therefore interpreted as a variety of that species.
While it shares some characters, like the checkered
Gamboa juniper; cedro, tscate (Spanish); tzotzil ni bark (not present in all varieties of J. deppeana), it
(Mexico) also has some distinct ones. The analysis presented
is far from convincing, with largely unresolved
Description relationships among the varieties of J. deppeana,
but high statistical support for a clade with J. gam-
Shrubs or small trees to 812 m, monoecious or boana and J. deppeana var. robusta. If J. gamboana
dioecious; trunk very short or straight, to 5090cm were indeed most closely related to J. deppeana var
diam. Bark on trunks fissured, 1015mm thick robusta, the geographical distribution of these two
and hard, forming longitudinal plates, lower part taxa would be hard to explain, as they are separated
of trunk often with quadrangular plates, weather- by populations of J. deppeana var. deppeana. A bet-
ing grey. Branches spreading or ascending, forming ter resolved cladogram based on a wider sampling
a broadly conical or pyramidal crown, in old trees of data is needed to justify this new taxonomy and J.
becoming irregular and spreading. Foliage branches gamboana is therefore here maintained as a distinct
spreading or drooping, ultimate branchlets spread- species.
Distribution irregularly disposed at angles between 2545
degrees, mostly shorter than 20mm, slender or
Mexico: Chiapas; Guatemala: Huehuetenango. thicker in one variety, 0.91.5mm diam., more
TDWG codes: 79 MXT-CI 80 GUA or less quadrangular in ultimate branchlets, per-
sistent. Leaves decussate, decurrent, scale-like,
Ecology imbricate, with free apex on ultimate branchlets,
1.52 0.61mm, rhombic to rhombic-lanceolate;
This species occurs in the understorey of, or mixed margins entire, hyaline; apex obtuse, mucronate
with other trees in Quercus-Pinus-Juniperus forests, or acute-acuminate; stomata few at base abaxially,
bearing many epiphytes indicating frequent fog con- in one concave field adaxially; abaxial gland more
ditions, on often rocky soil types; also in open pine or less conspicuous in basal to central part of leaf, 433
forest, e.g. with Pinus oocarpa, where J. gamboana rounded to oval; leaf colour green. Pollen cones
attains greatest size. The known altitudinal range is terminal on ultimate branchlets, subglobose, very
18202200 m a.s.l. small ca. 2mm; microsporophylls 68, decussate,
peltate, with rounded and hyaline upper margin,
Conservation abaxially bearing 23(4) globose pollen sacs. Seed
cones terminal on straight, short ultimate branch-
Due to its very limited range (most collections lets, maturing in one season, becoming globose
known are from Chiapas, Mexico) in an area which (with a single seed) or broad pyriform to nearly
suffers rapid deforestation, the species is likely to reniform, 45(6) mm wide, usually soft pulpy,
have suffered a significant reduction in its area of resinous, reddish brown with glaucous-blue bloom.
occupancy (AOO) as well as its overall numbers of Bract-scale complexes 4, decussate, the lower pair
mature trees. The situation in Guatemala with this small and sterile, entirely fused; sutures or bract
species is poorly known. tips invisible. Seeds 12 per cone, ovoid-globose,
IUCN: EN [B2ab (ii, iii, v)] 23mm diam., light brown.
Distribution
Juniperus gracilior Pilg., in Urban, Symb. Antill. 7:
481. 1913. Hispaniola (Dominican Republic and Haiti), very
localized in relict populations.
Etymology TDWG codes: 81 DOM HAI-HA
Juniperus gracilior Pilg. var. gracilior. Type: Trees. Ultimate branchlets spreading at 3045
Dominican Republic: Cordillera Central, Sierra degrees, slender, 0.91.0mm diam., more or less
434 de la Constanza, Las Caitas, M. Fuentes & quadrangular. Leaves slightly longer than broad;
D. Goodwin 1939 (lectotype NY). apex acuminate or mucronate; gland conspicuous,
near base.
Description
Distribution
Trees. Ultimate branchlets drooping or pendulous,
spreading at 2530 degrees, slender, 0.91.1mm Hispaniola: Haiti, Massif de la Selle (Morne la Selle,
diam., more or less quadrangular. Leaves distinctly Morne la Visite?); Dominican Republic (Sierra
longer than wide, obtuse or acute-acuminate, gland Baoruco).
central. TDWG codes: 81 DOM HAI-HA
Distribution Conservation
Hispaniola: Dominican Republic (La Vega Prov., This variety is now limited to two or perhaps three
near and 14 km W of Constanza; Azua Prov., Valle localities, where very few trees remain.
del Yaque). IUCN: CR (D)
TDWG codes: 81 DOM
Distribution Etymology
Boreal and Subarctic North America, from Alaska The species epithet denotes the country of origin,
(disjunct) to Newfoundland, more scattered further i.e. India, from where it was first described.
south in the USA.
TDWG codes: 70 ASK NUN NWT YUK 71 ABT BRC Vernacular names
MAN SAS 72 LAB NBR NSC NFL-NE ONT PEI QUE
436 73 MNT WYO 74 ILL IOW MIN MSO NEB NDA SDA Black juniper, Wallichs juniper; dian zang fang zhi
WIS 75 MAI MAS MIC NWY VER 78 VRG bai (Chinese)
Ecology Description
This species occurs in a variety of habitats, usually (Decumbent) shrubs, rarely small trees to 15(20) m
on more or less open ground; on sandy beaches tall, dioecious; multistemmed or monopodial; trunk
and in sand dunes, on dry rocky slopes and out- up to 1.2 m d.b.h. Bark on larger stems exfoliating in
crops, limestone ridges, in dry barrens, in grassland sheets or longitudinal strips, weathering dull brown.
(prairies), open bogs or muskeg (Picea mariana / Branches ascending or spreading, foliage branches
Pinus banksiana bog-woodland), or heathland (e.g. dense, short, stiff and spreading or erect, forming
Empetrum nigrum), or on stream banks, often form- a dense, broad pyramidal to eventually rounded
ing wide patches. The altitudinal range is from 10 m or irregular crown. Foliage branchlets ultimately
to 1160 m a.s.l. stout, quadrangular, sometimes more or less terete,
1.21.5mm wide, covered with appressed or some-
Conservation times spreading leaves, persistent. Leaves on mature
plants eventually scale-like, decussate, sometimes
IUCN: LC 3-whorled, imbricate, decurrent, 1.22 11.2mm,
triangular-rhombic, obtuse; margins entire; scale
Uses leaves amphistomatic, stomata in 2 inconspicuous
lines on each side mostly near base; glands central
Creeping juniper is a desirable species for horticulture in a groove, elliptic or oblong, sometimes absent;
in cool temperate to cold regions, since 1830 known in leaf colour green or slightly glaucous green. Pollen
cultivation in England. There are now numerous gar- cones numerous, solitary, terminal on short branch-
den forms (cultivars) known, selected both in Europe lets, subglobose to ovoid-globose, 23mm long;
and North America and widely used in gardens and microsporophylls 68, decussate, peltate-cordate,
parks. It makes an excellent ground cover on sandy with rounded entire hyaline margins, bearing 23
soils and in rockeries. Many forms that originated abaxial pollen sacs. Seed cones terminal on short
in cultivation have in the past been given botanical erect branchlets, maturing in the second season to
latinized names but since they are not taxa but culti- subglobose or ovoid, (4.5)513 48mm, lustrous
vars these names should be changed to cultivar names blue-black or brownish black, soft cones. Bract-
written in inverted commas (or preceded by cv.), scale complexes 36, decussate or (2) 3-whorled,
capitalized and in roman type; e.g. Juniperus horizon- entirely fused, sometimes incompletely covering
talis var. viridis Grootendorst becomes J. horizonta- the seed; bract tip subapical, small, ca. 0.5mm; sur-
lis Viridis. All such names were excluded from my face smooth; scale tissue succulent, resinous. Seeds
Monograph of Cupressaceae and Sciadopitys (Farjon, single, ovoid, laterally compressed or more or less
2005a) which only includes taxa (ranked entities clas- flattened, 56 4mm, shallowly grooved, pale yel-
sifying variation that occurs in nature). lowish brown.
Distribution Juniperus indica Bertol. var. indica. Type:
Illustration in Bertoloni, Misc. Bot. 23: 16, t. 1. 1862
Himalaya, E into high mountains of China. Bhutan; (lectotype).
China: W Sichuan, S Xizang [Tibet], NW Yunnan;
India: Himachal Pradesh, Uttar Pradesh, Kashmir; Juniperus wallichiana Hook. f. & Thomson ex
Nepal; N Pakistan; Sikkim. E. Brandis, Forest Fl. N.W. & Central India: 537. 1874.
TDWG codes: 36 CHC-SC CHC-YN CHT 40
EHM-BH EHM-DJ EHM-SI NEP PAK WHM-HP Description
WHM-JK WHM-UT
Erect shrubs to small trees to 15(20) m tall. Foliage
Ecology branches spreading or more or less erect, dense with 437
short branchlets. Seed cones when mature mostly
From upper montane coniferous forest and wood- (broadly) ovoid, 813 58mm, blue-black or
land in pure stands, or with e.g. Abies, Pinus, brownish black.
Cupressus torulosa, or in Betula utilis subalpine
woodland, to alpine heath and grassland and into Distribution
the bare moraines and scree of the niveous zone. The
altitudinal range is from 3600 m to 4800 m a.s.l. As As the species.
an understorey shrub or tree in coniferous forest it
is often accompanied by J. squamata, Rhododendron Conservation
spp., Rosa, and Cotoneaster. Above the tree line it
can form juniper-rhododendron thickets, grow in IUCN: LC
Kobresia-Stipa turf with dwarfed alpine shrubs (e.g.
Rhododendron, Salix, Juniperus squamata), or occur
scattered on moraines and consolidated scree slopes Juniperus indica Bertol. var. caespitosa Farjon,
of granite or gneiss or other metamorphic acidic Monogr. Cupressaceae & Sciadopitys: 313. 2005.
rock, at the highest altitudes exclusively on S-facing Type: Nepal: Dhaulagiri Himal, Dolpo, Mugu
slopes. The climate is high montane to alpine with a Karnali, S of Mugu, S. Miehe 9903001 (holotype K).
pronounced monsoon phase delivering heavy pre-
cipitation (much as snow) from May to October. Description
Juniperus jaliscana Martnez, Anales Inst. Biol. Mexico: S Durango, NW Jalisco, only known from
438 Univ. Nac. Mxico 17 (1): 69. 1946. Type: Mexico: two localities.
Jalisco, Talpa de Allende, Cumbre Blanca, TDWG codes: 79 MXE-DU MXS-JA
M. Martnez & A. Q. Gonzalez 7002 (holotype
MEXU). Ecology
plate 17. Juniperus monosperma. 1. Habit of shrub. 2. Branch with foliage. 3. Branchlet with leaves.
4. Leaves with gland. 5. Pollen cone. 6, 7. Seed cones, one opened to show seed. 8. Seeds.
few stomata abaxially on lower margin, stomata from Ficus religiosa forest, with an understorey of
scattered adaxially, covered by thick cuticular wax; Ribes, Rosa and Rubus (herbarium collection Zanoni
glands central, elliptic or nearly linear, or absent on 2718, Mexico, Federal District, 2895 m). The decum-
many smallest leaves; leaf colour light to dark green bent shrub form is common at or above the tree line
or glaucous green. Pollen cones terminal, solitary, (usually with Pinus hartwegii) on ridges of basalt or
oblong, 34 22.5mm, more or less quadrangular consolidated scree, with grasses e.g. Calamagrostis
in cross section; microsporophylls 1012, decussate, and Festuca, shrubs such as Ribes, and some alpine
peltate, often keeled, bearing 34 abaxial pollen sacs. herbs, or on rocks and cliffs to the limit of vegeta-
Seed cones terminal on very short branchlets; mature tion on Mexicos highest volcanoes. The altitudinal
cones globose or subglobose, 69(10) mm diam., range of the species is 20004270 m a.s.l. Despite
442 lustrous, purplish black or purplish red, sometimes high precipitation on these mountains, the environ-
dull orange-brown, usually with a glaucous bloom. ment is decidedly xeric due to the porosity of volca-
Bract-scale complexes 6, completely fused, forming a nic substrates, but snowmelt may provide a relatively
soft pulpy, succulent, resinous cone with smooth sur- reliable, slow-release moisture source in spring and
face and minutely exserted (lower) bract tips. Seeds early summer.
(2)37(9) per cone, angular ovoid, irregular but
more or less flattened, 35 23mm, small relative Conservation
to cone size, with a small light hilum at or near base,
grooved or pitted proximally, with resin in pits, lus- IUCN: LC
trous (dark) chestnut-brown.
Uses
Taxonomic notes
This species is not in cultivation, but the decumbent
Recently, Adams (op. cit.) raised Juniperus monticola alpine form would probably be suitable for rock gar-
var. monticola f. compacta to the rank of species: J. dens and withstand low winter temperatures.
compacta (Martnez) R. P. Adams, based on analysis
of DNA sequence data and biochemistry. No mor-
phological characters seem to separate J. monticola Juniperus occidentalis Hook., Fl. Bor. Amer. 2 (10):
from J. compacta other than the habit mentioned by 166. 1838.
Martnez in his original description. Whilst it may be
so that these biochemical differences exist (and even Etymology
could turn out to be consistent with wider sampling),
it remains to be demonstrated that it is taxonomically The Latin species eithet means from the west.
informative to base species concepts on such data,
creating, as Adams admits, cryptic species. Vernacular names
W USA: California, W Idaho, Nevada, Oregon, Juniperus occidentalis Hook. var. occidenta-
Washington. lis. Type: USA: Washington, Columbia River,
TDWG codes: 73 IDA ORE WAS 76 CAL NEV [Common on the higher parts of the Columbia, at
the base of the Rocky Mountains], D. Douglas s.n.
Ecology (holotype K).
Description Conservation
N Algeria; also reported from E Portugal and central Juniperus navicularis Gand., Bull. Soc. Bot. Prance
Spain. 57: 55. 1910.
TDWG codes: 12 POR SPA-SP 20 ALG
Description
Conservation
Shrubs to 2 m with fastigiate branching; foliage
IUCN: LC branchlets spreading. Leaves spreading at 7090
degrees from shoot, (4)612 1.1.5mm, boat rhombic, gibbous; margins entire or minutely den-
shaped (but on whip shoots longer and linear-acic- ticulate; apex obtuse, sometimes acutish; stomata in
ular), subobtuse to acute (pungent on whip shoots). two small groups abaxially near base, adaxially in
Pollen cones 23mm long, ovoid. Seed cones sub- two narrow bands; glands conspicuous, sometimes
globose to ovoid, 710mm long, yellowish or red- inconspicuous, central, oval to linear, usually active;
dish but not glaucous or pruinose when growing, leaf colour lustrous light green or greyish green.
red when ripe. Pollen cones numerous, terminal on short ultimate
branchlets, ovoid or short cylindrical, 24mm long;
Distribution microsporophylls 616, decussate or sometimes
ternate, peltate, with minutely denticulate margins,
448 SW Portugal, adjacent part of Spain. bearing 35 abaxial pollen sacs. Seed cones termi-
TDWG codes: 12 POR SPA-SP nal on short branchlets, growing in two seasons to
ochraceous, red-brown or purplish black, sometimes
Conservation pruinose, (sub)globose to ovoid, (5)712(14) mm
diam., soft pulpy or fibrous, maturing soft succulent.
IUCN: NT Bract-scale complexes 46(9), decussate or ternate,
entirely fused, smooth, lustrous or slightly rugose
when dry, with triangular bract apices protruding
Juniperus phoenicea L., Sp. Pl. 2: 1040. 1753. from transverse ridges (sutures). Seeds 38 per cone,
relatively small, unequal to 4mm long, triangular or
Etymology flattened.
IUCN: LC
Etymology ridges, with resin in the grooves or pits, light yellow-
ish brown to chestnut brown, with a large, lighter
This species was named after Gifford Pinchot, Chief bilobed hilum.
Forester under US President Theodore Rooseveldt
and an early conservationist. Distribution
Procumbent or erect shrubs to (small) trees, monoe- This species occurs from subalpine coniferous for-
cious, to 10 m but reputedly attaining 30 m. Bark on ests with Abies, Larix, Picea and Pinus to beyond the
large stems fissured, exfoliating in long strips, weath- tree line in alpine meadows and high steppes to near
ering grey-brown. Branches spreading or ascending, the line of perennial snow or glaciers, at altitudes
very dense in procumbent shrubs, forming a spread- between 2650 m and 4850 m a.s.l.
ing or rounded crown. Foliage branches short, pat-
ent, drooping or downcurved, or pendulous, ultimate Uses
branchlets in shrubs 1030mm long, in trees with
pendulous branchlets much longer, stout or slender, The subalpine to alpine decumbent shrub forms are 451
subterete to more or less 6-angled, sometimes quad- excellent plants for cultivation in cool temperate to
rangular, covered with imbricate scale leaves. Leaves cold climate regions. A few clones are in cultivation
in alternate whorls of 3, rarely decussate, imbricate, in the West but are rare; these are commonly J. pingii
decurrent at base, appressed with incurved apices or var. wilsonii. The tree form (var. pingii) is occasion-
sometimes curved but free, lanceolate (naviculate) ally seen in cultivation in China; in Europe some
to broadly subulate, free portion broadest below introductions are likely to masquerade under the
middle section, incurved, (2)35(7) 11.5mm, names J. recurva and/or J. squamata which are actu-
concavo-convex; abaxial surface distinctly keeled or ally this species. Several selected cultivars are in the
ridged, especially towards the apex, smooth, glau- horticultural trade.
cous green; adaxial surface with two whitish bands
proximally separated by a flat or slightly raised 4 varieties are recognized:
midrib; margins entire; apex acuminate to pun-
gent; epistomatic, stomata in two converging bands.
Pollen cones axillary on very short dwarfed shoots, Juniperus pingii W. C. Cheng var. pingii. Type:
solitary, globose, 34mm diam.; microsporophylls Illustration in Y. de Ferr, Bull. Soc. Hist. Nat.
69, ternate, rarely decussate, more or less cordate, Toulouse 79: 75. 1944 (lectotype).
obtuse or cuspidate, bearing 23 abaxial globose pol-
len sacs. Seed cones axillary on very short dwarfed Description
shoots, growing to globose or ovoid cones (5)69
56mm in two seasons, ripening to lustrous pur- Arborescent shrubs to trees possibly to 30 m tall.
plish black or bluish black. Bract-scale complexes 3, Foliage branches drooping or pendulous; ultimate
rarely 4, entirely fused, not discernible in soft suc- branchlets usually 6-angled, or subterete, slender;
culent-pulpy mature cones except for the presence leaves nearly straight on young trees, later curved.
of 3(4) subapical minute bract tips. Seeds ovoid-
globose, 56 4mm, with several shallow, resinous Distribution
grooves or pits towards base.
China: W Sichuan, NW Yunnan, SE Xizang [Tibet].
Distribution TDWG codes: 36 CHC-SC CHC-YN CHT
Distribution
Juniperus pingii W. C. Cheng var. wilsonii (Rehd.)
China: NW Yunnan (imperfectly known from the Silba, Phytologia Mem. 7: 36. 1984. Juniperus
type collection near Zhongdian only). squamata Buch.-Ham. ex D. Don f. wilsonii Rehd.,
TDWG codes: 36 CHC-YN J. Arnold Arbor. 1: 191. 1920. Type: China: Sichuan,
W Sichuan, E. H. Wilson 985 (lectotype E).
Conservation
Description
The distribution of this form seems to be much more
limited than that of the species as a whole, but its Juniperus pingii W. C. Cheng var. carinata Y. F. Yu &
extent remains insufficiently known. It was described L. K. Fu, Novon 7: 443. 1998; Juniperus carinata (Y.
as a small tree; the most common and widespread F. Yu & L. K. Fu) R. P. Adams, Biochem. Syst. Ecol.
variety, J. pingii var. wilsonii (Rehd.) Silba is a low 28: 541. 2000.
shrub. Procumbent to erect shrubs or arborescent shrubs
IUCN: DD to 6 m tall. Foliage branches not pendulous, usually
curved, stout, subterete or prominently 6-angled.
Leaves appressed or sometimes curved but free,
Juniperus pingii W. C. Cheng var. miehei Farjon, slightly to strongly incurved.
Monogr. Cupressaceae & Sciadopitys: 346. 2005.
Type: China: Xizang (Tibet), Zangbo River, Upper Taxonomic notes
Zangbo basin, S of Saga, along road to Gyirong,
G. Miehe & S. Miehe 953201 (holotype K). The distinction between Juniperus pingii var. wilso-
nii and Juniperus squamata has been a contentious
Description subject for some time. Rehder (op. cit.) originally
described this taxon as only a form of J. squamata,
Procumbent shrubs with very dense, short foliage. but it was treated as a variety of J. squamata in sub-
Ultimate branchlets predominantly quadrangular, sequent arboricultural and floristic literature, and
with decussate, short leaves. Seed cones globose, as a cultivar name under J. squamata in the fourth
57mm diam. edition of Dallimore & Jacksons Handbook of
Coniferae and Ginkgo (4th ed., 1966). It was trans-
ferred to Sabina pingii var. wilsonii in FRPS 7 (Cheng
& Fu, 1978); this became Juniperus pingii var. wilsonii Description
in Flora of China 4 (1999) and again Sabina pingii var.
wilsonii in Higher Plants of China 3 (Fu et al., 2000). Trees to 3040 m tall, monoecious or dioecious;
Its morphological differences with J. squamata are trunk to 1.52 m d.b.h. Bark on larger stems fissured,
mainly in the leaves, which are more similar to those exfoliating in long, narrow, fibrous strips, (pale)
of J. recurva. In high altitude subalpine meadows brown weathering grey-brown. Branches long,
(40004300 m) of Baima Shan, NW Yunnan J. pingii spreading or ascending, foliage branches drooping
var. wilsonii and J. squamata grow together, some- or pendulous, lax, forming a pyramidal crown in
times accompanied by a form that could be a hybrid young trees, but later broader, rounded and finally
of the two (pers. obs., September 2000). Otherwise flat-topped or open and irregular in old trees. Foliage
the two taxa are distinct in this area. branchlets ultimately slender, (weakly) quadrangu- 453
lar, 0.61mm wide, covered with closely appressed
Distribution leaves, persistent. Leaves on mature plants scale-
like, decussate, imbricate, decurrent, on ultimate
China: S Gansu, NW Hubei, S Qinghai, S Shaanxi, branchlets 0.51 0.40.6mm, triangular-rhombic,
Sichuan, NW Yunnan, Xizang [Tibet]. acute; margins entire; stomata in 2 inconspicuous
TDWG codes: 36 CHC-HU CHC-SC CHC-YN lines on each side mostly near base; leaves abaxi-
CHN-GS CHN-SA CHQ CHT ally glandular; gland central, conspicuous, elliptic
or nearly linear; leaf colour yellowish green or light
Ecology green. Pollen cones numerous, solitary, terminal
on short branchlets, subglobose to ovoid, 35
A high altitude variety commonly replacing subal- 23mm; microsporophylls 1012, decussate, pel-
pine coniferous forest or woodland above the tree tate, with minutely denticulate margins, bearing 23
line, forming thickets or patchy mats of decumbent abaxial pollen sacs near lower margin. Seed cones
juniper among scree, rocks, or in alpine grassland or terminal on short, erect branchlets, maturing in
steppe. The altitudinal range is 26504800 m a.s.l. the second season to (sub)globose, 37mm diam.,
brown, blue or purplish black, usually glaucous or
Conservation pruinose. Bract-scale complexes 4(6), decussate,
entirely fused, bract tip small, ca. 0.3mm, scale tis-
IUCN: LC sue dry pulpy. Seeds (1)23(4) per cone, angu-
lar-ovoid, more or less flattened on one side, 45
33.5mm, yellowish brown, with lighter hilum
Juniperus procera Hochst. ex Endl., Syn. Conif.: 26. at base.
1847. Type: Ethiopia: Gonder Prov., Amhara, Adda
Mariam, near Enschedcap [ad ecclesiam Adda Distribution
Mariam prope Enschedcap], G. H. W. Schimper
537 (lectotype L). NE, E & S Tropical Africa: Congo Republic, Djibouti,
Eritrea, Ethiopia, Kenya, Malawi, Somalia, Sudan
Etymology (near Red Sea), Tanzania, Uganda, NE Zimbabwe;
Arabian Peninsula: Saudi Arabia (Asir Range),
The species epithet means slender or tall. Yemen.
TDWG codes: 23 ZAI 24 DJI ERI ETH SOM SUD 25
Vernacular names KEN TAN UGA 26 MLW ZIM 35 SAU YEM-NY
Distribution Description
Japan: Kyushu, Nansei-Shoto [Ryukyu Islands] Trees, rarely shrubs, to 15(20) m, monoecious;
(Okinawa Group, Sakishima Group). trunk up to 1 m d.b.h. Bark on larger stems exfoliat-
TDWG codes: 38 JAP-KY NNS OGA ing in longitudinal strips, weathering whitish grey or
grey-brown. Branches spreading, smaller branches
Ecology drooping to subpendulous, forming a broadly 455
rounded or irregular crown. Foliage branchlets
The ecology of this species is poorly known. It occurs slightly curved or drooping to pendulous, quadran-
at high altitudes near or on island mountain sum- gular or subterete, 1.21.5mm wide if covered with
mits [Ohwi, Fl. Japan: 118 (1965) has it on seashores appressed scale leaves, often with transitional leaves
[of] Kyushu but this is J. rigida subsp. conferta], or juvenile leaves, appearing ragged, persistent.
where it forms decumbent mats over rocks. Leaves on mature plants scale-like as well as acicular
(scale leaves predominant on old trees), decussate,
Conservation or 3-whorled, imbricate, decurrent, scale leaves on
ultimate branchlets 1.52.5(3) 11.5mm, triangu-
IUCN: LC lar-rhombic, with free apex, acute, transitional and
juvenile leaves with spreading parts 48(10) 1.5
Uses 1.7mm; margins entire, pungent; scale leaves weakly
amphistomatic, juvenile and transitional leaves
This species is popular in gardens as a ground cover- epistomatic; stomata in two bands separated by an
ing evergreen conifer shrub and a limited number inconspicuous midrib; scale leaves abaxially glandu-
of cultivars are in the trade. It is used more often lar; gland near base, elliptic to orbicular, convex; leaf
in Japan than in Europe, although it is not uncom- colour green or glaucous green. Pollen cones solitary,
mon there. Unlike some other species with a similar terminal on short branchlets, subglobose to ovoid-
growth habit, Creeping juniper faithfully follows globose, 23mm long; microsporophylls 68(10),
the contours of rock gardens without erecting itself decussate, peltate-cordate, with entire hyaline mar-
or massing in one spot. It is therefore very effective gins, bearing 23 abaxial pollen sacs near lower mar-
in such gardens, especially in traditional oriental gin. Seed cones terminal on short erect branchlets,
gardens. maturing in the second season to become ovoid,
(8)1013 910mm, lustrous blue-black or pur-
plish black, slightly pruinose and soft. Bract-scale
Juniperus przewalskii Kom., Bot. Mater. Gerb. complexes 6, decussate or (2) 3-whorled, entirely
Glavn. Bot. Sada RSFSR 5: 28. 1924. Type: China: fused; bract tips usually hidden or minute; surface
Gansu, Qilian Shan, Daban Shan, [jugum a fluv. smooth; scale tissue succulent, resinous. Seeds 1(2)
Tetung versus in regione alpina sol. humido, 13.000 per cone, ovoid-globose, laterally compressed or
ft. alt.], N. M. Przewalski 318 (lectotype LE). slightly flattened, (7)812 (6)710mm, shal-
lowly or deeply grooved, with resin pits, light brown.
Etymology
Distribution
This species name commemorates the Polish soldier,
traveller and naturalist Nikolai M. Przewalski (1839 China: SW Gansu, E Qinghai, NW Sichuan (upper
1888), who explored Central Asia and (greater) tributaries of the Huang He).
Mongolia. TDWG codes: 36 CHC-SC CHN-GS CHQ
Ecology Vernacular names
Juniperus przewalskii occurs in the ecotone between Xinjiang juniper; xin jiang fang zhi bai, ka shi fang
subalpine forest to alpine meadows or steppe, as well zhi bai, kun lun fang zhi bai (Chinese); no local
as in Juniper-Picea crassifolia woodland, on steep, (Kyrgiz) names have been recorded.
usually S-facing, calcareous slopes, with ground
cover of grasses (e.g. Stipa splendens), shrubs (e.g. Description
Salix sp.) or perennial herbs (e.g. Polygonum
viviparum), on dry or moist substrates. The altitude (Decumbent) shrubs or small trees to 810 m, dioe-
ranges between (1900)2250 m and 4000(4500) m cious, sometimes monoecious; multistemmed or
456 a.s.l. The climate is cold temperate with substantial monopodial, with a short trunk up to 1 m d.b.h. Bark
periods of snow cover in winter. on larger stems shaggy, exfoliating in short strips
remaining partly attached, weathering grey-brown.
Conservation Branches ascending, assurgent or spreading, smaller
branches numerous, short, stiff and spreading, form-
The relatively restricted distribution and its occur- ing a dense, spreading or irregular crown. Foliage
rence in a region where forest cover is scattered and branchlets thick, quadrangular, sometimes more or
limited to suitable slopes make it potentially vulner- less terete, 1.52mm wide, covered with appressed
able to logging and/or deforestation resulting from or sometimes apically free leaves, persistent. Leaves
increased human pressure on these resources. As on mature plants normally scale-like, decussate,
yet, however, there is no immediate concern regard- imbricate, decurrent, 1.32 11.2mm, triangular-
ing this species. rhombic, gibbous, obtuse, on older branchlets to 4
IUCN: LC 1.5mm, mostly appressed but several with free apex;
margins entire or sometimes minutely and intermit-
Uses tently denticulate; stomata in 2 conspicuous short
bands on each side mostly near base; scale leaves
No uses have been recorded, but this species is likely abaxially glandular; gland central in a groove, ellip-
to be used for firewood as it is one of the few trees tic or oblong, sometimes absent; leaf colour green or
that exists in some parts of its range. yellowish green, sometimes slightly glaucous green.
Pollen cones numerous, solitary, terminal on short
branchlets, subglobose to globose, 23mm long;
Juniperus pseudosabina Fisch. & C. A. Mey., Index microsporophylls 68, decussate, peltate-cordate,
Sem. Hort. Petrop. 8 (1841): 24, 65. 1842. Type: with rounded entire hyaline margins, bearing 23
Kazakhstan: Dzhungarskiy Alatau, Tarbagatay Mts., abaxial pollen sacs. Seed cones terminal on short
A. G. von Schrenk s.n. (lectotype K). Fig. 144, 145 erect branchlets, maturing in the second season to
become ovoid, (7)814 (6)710mm, lustrous
Juniperus turkestanica Kom., Bot. Mater. Gerb. blue-black or purplish black and soft. Bract-scale
Glavn. Bot. Sada RSFSR 5: 26. 1924; Juniperus pseu- complexes 46, decussate, entirely fused; bract tip
dosabina Fisch. & C. A. Mey. var. turkestanica (Kom.) ca. 0.5mm; surface smooth; scale tissue succulent,
Silba, Phytologia Mem. 7: 36. 1984. resinous. Seeds 1 per cone, ovoid-ellipsoid, laterally
compressed, 68 4.56.5mm, shallowly grooved,
Etymology light brown.
The species epithet means similar but not equal Taxonomic notes
to (J.) sabina, a species that occurs in the same
region. Russian botanists and ecologists treat the taller
shrub in the west as a distinct species J. turkestanica
Kom. No botanical characters appear to separate the
two and J. turkestanica is not a species, but only an
ecotype that predominates in the juniper forest and an omission that ought to be rectified. The species
woodland, an ecosystem not present in the eastern occurs in a continental climate at high elevations in
parts of the Central Asian mountains. mountains similar to the Alps and should be hardy
even in high latitudes.
Distribution
Central Asia: Afghanistan (Takhar), China Juniperus recurva Buch.-Ham. ex D. Don, Prodr. Fl.
(Xinjiang), Kazakhstan (southern mountains), Nepal. (2): 55. 1825.
Kirgyzstan, Mongolia, Pakistan (Baltistan, Hindu
Kush, Karakoram Range), India (Kashmir), Etymology
Tadjikistan, Uzbekistan (Turkestan Range). 457
TDWG codes: 32 KAZ KGZ TZK UZB 34 AFG 36 The species epithet refers to the (re)curved adult
CHX WHM-JK 37 MON 40 PAK type leaves.
Occurring in subalpine conifer forest with Picea Drooping juniper, Sacred juniper, Coffin juniper
schrenkiana, Pinus sibirica, or P. wallichiana, in (var. coxii); Pama (Hindi); chui zhi bai, xiao guo chui
juniper woodland (J. semiglobosa) and in mon- zhi bai (var. coxii) (Chinese)
tane to subalpine scrubland and steppe (pre-
dominantly Seriphidium maritimum [Artemisia Description
maritima] steppe rich in grasses and geophytes),
with e.g. J. sabina, Cotoneaster, Kobresia capillifo- Shrubs or trees to 15(40) m tall, monoecious, occa-
lia, Rhododendron anthopogon, Rosa, and Salix. The sionally dioecious; multistemmed or monopodial;
altitudinal range is from 1950 m to 4100 m a.s.l. It trunk up to 2 m d.b.h. Bark on larger stems exfo-
is often restricted to rocky outcrops and S-facing liating in longitudinal, curling strips, red-brown
slopes, especially in forests, and occurs on various weathering grey-brown. Branches ascending or
rock types and soil types from coarse gravel terraces spreading, higher order branches assurgent and
to dry S-exposed loess slopes. The climate is extreme drooping or very long and pendulous, forming a
continental with short, hot, dry summers and long, broad pyramidal to eventually rounded or irregular
cold, snowy winters. Both climatic factors and (pos- crown. Foliage branchlets numerous, crowded, lax,
sibly) grazing pressures are responsible for a shift more or less terete, covered with incurved or slightly
from erect to decumbent shrubs from west to east in spreading leaves, persistent. Leaves in alternating
its range. whorls of 3 (on whip shoots occasionally decussate),
imbricate, decurrent, mature type leaves 210 0.8
Conservation 1(1.2) mm, lanceolate-acicular, weakly keeled; mar-
gins entire, acute to pungent or acuminate, incurved
IUCN: LC or straight with a free apex; stomata abaxially in 2
inconspicuous lines on each side mostly near base,
Uses in two adaxial white primary bands separated by
an obscure midrib; leaf colour green or olive-green.
This species is more often shrubby than a tree and Intermediate leaf shapes, as well as juvenile-type
consequently it is less often used for firewood or leaves, usually present in mature trees and retained
small timber. The large blue cones (berries) are soft in one variety. Pollen cones numerous, solitary, axil-
and probably edible, but no commercial use of them lary on dwarfed shoots, ovoid-oblong, 56 23mm
was spotted in local markets on a trip to Kirgyzstan long; microsporophylls 1016, decussate, more or
in August 2000. Especially the shrubby form of this less remote, peltate-cordate, with entire-hyaline to
species would make a very atractive juniper in cul- erose margins, bearing 3 abaxial pollen sacs. Seed
tivation but it seems to be absent in gardens; this is cones axillary on dwarfed shoots, maturing in the
458
plate 18. Juniperus recurva var. recurva. 1. Habit of tree. 2, 3. Branches with leaves and seed cones.
4. Branchlet with leaves. 5, 6. Leaves. 7. Branchlet with pollen cone. 8. Microsporophyll with open pollen
sacs and pollen. 9. Seed cone. 10. Seed.
second season becoming ovoid or subglobose, 612 Juniperus recurva Buch.-Ham. ex D. Don
59mm, lustrous purplish black and soft. Bract- var. recurva. Type: Nepal: Narainghet (?)
scale complexes 3, entirely fused; bract tip subapical, [Narainhetty], F. Buchanan-Hamilton s.n.
ca. 0.5mm; surface smooth; scale tissue succulent, (holotype BM). Pl. 18
resinous. Seeds 1 per cone, ovoid-globose, broadest
at base, 68 56mm, shallowly grooved with dark Description
brown resin pits near base, light brown.
Shrubs or small trees to 15 m tall, foliage branches
Distribution drooping, not pendulous. Leaves 23(5) 11.2mm,
sometimes a few branches with intermediate leaves
Himalaya: Nepal, Sikkim, Bhutan, India (Arunachal slightly longer, incurved. Seed cones 712 69mm; 459
Pradesh, Assam); Myanmar [Burma]; China: S seeds 68 56mm.
Xizang [Tibet], NW Yunnan, SW Sichuan.
TDWG codes: 36 CHC-SC CHC-YN CHT 40 ASS-AS Distribution
EHM-AP EHM-BH EHM-DJ EHM-SI NEP WHM-UT
41 MYA As for the species.
Ecology Ecology
In montane evergreen rainforest (var. coxii) ascend- In high montane to subalpine coniferous forest,
ing to and beyond the tree line in alpine scrub and with Abies spp. or Picea spp. and an understorey
rocky meadows (var. recurva). The altitudinal range of Rhododendron, in Rhododendron thickets in the
is 24004500 m a.s.l. See for more detailed accounts ecotone between forest and alpine meadows, and
under the varieties. in the latter in mixed shrub communities with
e.g. Rhododendron, Salix, Cotoneaster, Berberis,
Uses Lonicera, Spiraea, and Potentilla. The altitudinal
range is 25004500 m a.s.l. It is common in rocky
In its native countries this species is used for timber areas (e.g. moraines) or in alpine meadows strewn
as well as ornamental trees in gardens of monaster- with boulders, usually on siliceous rock. In western
ies and temples. The wood and foliage are burned for Yunnan I observed that this variety is a pioneer after
incense in Buddhist temples. In Myanmar [Burma] destructive forest fires that had killed most trees of
the wood of large trees is used to make coffins. Abies and Tsuga in the area. It was not clear if, but
The drooping form J. recurva var. coxii is a highly it could be possible that the extensive stands of J.
ornamental tree much valued and often planted in recurva var. recurva observed in this area (Nu Shan,
regions of Europe with a mild, moist climate. Several NW of Caojian) are the result of past forest destruc-
cultivars have been selected, some with even more tion. The climate is more alpine with much longer
pendulous foliage than var. coxii. Decumbent shrubs periods of deep snow cover with this variety than
in cultivation under the name J. recurva (unless cul- with var. coxii.
tivars from proven J. recurva provenance) are proba-
bly J. pingii var. wilsonii; J. recurva is not a decumbent Conservation
shrub in the wild, but is often multi-stemmed. It is
one of the few junipers that require a good amount IUCN: LC
of rainfall and thrive best in a maritime, cool but
mild climate.
2 subspecies are recognized: Juniperus rigida Siebold & Zucc. subsp. litoralis
Urussov, Bjull. Glavn. Bot. Sada 122: 56. 1981; Juniperus
rigida Siebold & Zucc. var. litoralis (Urussov)
Juniperus rigida Siebold & Zucc. subsp. rigida. Kozhevnikova, Fl. Ross. Dalnego Vostoka: 42. 2006.
Type: Japan: Honshu [in Japoniae ad monte
Hakone (Hakonegasaki?)], J. Brger, [comm. P. F. Vernacular names
von Siebold 1842] s.n. (lectotype M).
Shore juniper; Hai-nezu, Hai-muro, Hai-matzu
Description (Japanese)
This decumbent juniper is sometimes still treated as Usually decumbent, sometimes ascending or erect
a distinct species. Apart from the different growth shrubs, rarely a stunted tree to 1012 m with a short,
462 form, its leaves are larger (wider) than those in the leaning trunk to 2 m diam., dioecious or monoe-
erect growing form of the species. Ontogenetically, cious. Bark on branches smooth, soon flaking, yel-
this leads to larger ovuliferous cones and larger lowish brown, becoming grey-brown. Branches
seeds as well. As with the Mediterranean coastal spreading horizontally or assurgent, of higher order
subspecies J. oxycedrus subsp. macrocarpa, J. rigida mostly assurgent or erect, extending broom-like,
subsp. conferta is apparently a littoral variant that forming matting crowns in decumbent shrubs and
is ecologically separated from the remainder of dense, flat crowns in taller plants and small trees.
the species. The two subspecies have quite similar Foliage branchlets numerous, assurgent to erect,
morphological traits, which are perhaps of adap- slender or sometimes thicker 0.81(1.4) mm diam.,
tive value to the shifting dune sands on maritime subterete to nearly quadrangular in branchlets with
coasts. scale leaves, persistent. Leaves decussate, ocassion-
ally in alternating whorls of 3, short decurrent, of
Distribution two types: needle-like and, more commonly, scale-
like (acicular leaves in young plants and persisting
Japan: Hokkaido, Honshu, Kyushu; Russian Far East: in one variety). Needle-like leaves linear-subulate,
Sakhalin. (4)610 0.51mm, pungent, with a single or two
TDWG codes: 31 SAK 38 JAP-HK JAP-HN JAP-KY adaxial stomatal bands wider than green margins.
Scale leaves always decussate, imbricate, closely
Ecology appressed, on ultimate branchlets 12.5 0.61mm,
ovate-rhombic to rhombic-lanceolate, abaxially
This subspecies is strictly littoral and occurs on with a conspicuous central, elliptic, yellowish gland,
sandy ocean shores, forming dense, spreading mats dark lustrous green to yellowish green, strongly aro-
of shrubby vegetation covering old beaches and matic when bruised; margins entire; apex obtuse
dunes. Usually the vegetation is very open and has to acute; epistomatic or amphistomatic, stomata in
a pioneer character, but this juniper will sometimes two small bands. Pollen cones terminal on ultimate
occur (persist) in Pinus woodland near the shore. branchlets with scale leaves, 34mm long, ellipsoid;
microsporophylls 1016, decussate, peltate-cordate,
Conservation convex; upper margin erose, abaxially bearing 24
oblong pollen sacs. Seed cones terminal on recurved
IUCN: LC short shoots 310mm long with small scale leaves,
maturing in 12 years becoming (irregularly) glo-
bose, 48mm, usually soft pulpy, resinous, purplish
Juniperus sabina L., Sp. Pl. 2: 1039. 1753. brown, black-brown or blue. Bract-scale complexes
4, decussate, entirely fused, sutures partly visible as a
Etymology curved ridge terminating in a minute bract tip. Seeds
13(5) usually 2 per cone, ovoid-flattened, 35mm
The origin of this epithet is probably the French long, with proximal, shallow resin pits and longitu-
name of this species. dinal, shallow grooves, yellowish brown.
Distribution Juniperus sabina L. var. sabina. Type: [local-
ity unknown (Habitat in Italia, Sibiria, Olympo,
SW, Central & SE Europe, N Africa (Atlas Mts.), Ararat, Lusitania)], Herb. Burser XXV: 59
Ukraine: Krym [Crimea], Caucasus, Russia (Altai (lectotype UPS). Fig. 146
& Ural Mountains), Iran, Kazakhstan, Kirgyzstan,
Turkey, Mongolia, N & NW China. Sabina vulgaris Antoine var. yulinensis T. C. Chang
TDWG codes: 11 AUT-AU AUT-LI CZE-CZ CZE-SL & C. G. Chen, Acta Phytotax. Sin. 19 (2): 263. 1981;
GER POL SWI 12 FRA-FR SPA-SP 13 ALB BUL GRC Juniperus sabina L. var. yulinensis (T. C. Chang &
ITA-IT KRI ROM SIC-SI TUE YUG-CR YUG-MA 14 C. G. Chen) Y. F. Yu & L. K. Fu, Novon 7 (4): 444.
KRY RUE RUS UKR-MO 20 ALG MOR-MO 30 ALT 1998.
TVA WSB 32 KAZ KGZ TKM 33 NCS TCS 34 IRN TUR Juniperus sabina L. var. monosperma C. Y. Yang, Fl. 463
36 CHI-NM CHI-NX CHM-HJ CHN-GS CHN-SA Xinjiangensis 1: 305. 1993.
CHQ CHX 37 MON
Description
Ecology
Commonly decumbent shrubs, occasionally spread-
In montane to alpine vegetation formations, ranging ing shrubs or a small, stunted tree. Branches assur-
from alpine scree and rocks or meadows to nearly gent, erect or spreading. Leaves of two types: acicular
closed coniferous forest; in the east of its Asian range and scale-like; acicular leaves usually on seedlings
also in steppes and deserts. Altitudinal range 400 and young plants to 10 years old, merging to scale
3350 m a.s.l. More detailed accounts are given under leaves on most mature plants. Seed cones with
the varieties. (1)23(4) seeds, most frequently 2.
Uses Distribution
The decumbent shrub form has long been in culti- As for the species.
vation in Europe and is relatively common; several
cultivars (some with fastigiate growth habit) have Ecology
been named. Its cultivation is often more or less
restricted to countries where it is also native and In montane to subalpine coniferous forests of Larix,
where growers have experimented with this stock Picea and Pinus, gradually replacing these where
to produce cultivars; in other countries forms of the under human-imposed grazing regimes; also invad-
similar species J. chinensis seem to prevail. Some of ing into alpine meadows when old grazing patterns
the cultivars of J. sabina retain juvenile type (nee- are changed, e.g. intensified. In Central Asian moun-
dle) leaves, most have predominantly or exclusively tains it occurs on S-facing slopes in mountain pas-
scale leaves in mature plants. Forms with needle tures in a characteristic pattern of rounded patches,
leaves also occur in nature, so selection of this trait often mixed with J. pseudosabina. Its altitudinal
for horticulture is very easy where this variety (var. range is 7003000 m a.s.l. This subspecies is most
davurica) is available. The wood is of little value, but abundant on sunny, dry slopes in mountains with
was traditionally used in the Alps to make walking a mesic climate like the Alps; its drought tolerance
sticks. Oil is distilled from branches and foliage and accounts for its wider distribution in Asia into the
used for medicinal purposes; it has powerful diuretic Artemisia steppe and desert zones (var. arenaria). It
properties. is often found on limestone substrates but occurs on
granitic rock as well, especially on drier slopes.
3 varieties are recognized:
Conservation
IUCN: LC
Juniperus sabina L. var. arenaria (E. H. Wilson) Juniperus sabina L. var. davurica (Pall.) Farjon,
Farjon, Monogr. Cupressaceae & Sciadopitys: 366. Monogr. Cupressaceae & Sciadopitys: 367. 2005.
2005. Juniperus chinensis L. var. arenaria E. H. Juniperus davurica Pall., Fl. Ross. 1 (2): 13, t. 55.
Wilson, J. Arnold Arbor. 9: 20. 1928; Juniperus 1789. Type: Russia: Russian Far East, Amur River,
arenaria (E. H. Wilson) Florin, Acta Horti Berg. 14 [E Sibir. orientali], P. S. Pallas [ex herb. Pallas] s.n.
(8): 353. 1948. Type: China: Qinghai, Qinghai Lake, (lectotype BM).
J. F. Rock 13346 (holotype A).
Juniperus davurica Pall. subsp. maritima Urussov,
Juniperus sabina L. var. mongolensis R. P. Adams, Bjull. Glavn. Bot. Sada 122: 55. 1981.
Phytologia 88 (2): 182. 2006. Decumbent shrubs to 50cm. Branches ascending
464 to erect, whip shoots assurgent. Leaves of two types:
Description acicular and scale-like; acicular leaves usually present
from seedling to maturity, merging on some shoots
Decumbent or spreading shrubs to 1.5 m. Foliage to scale leaves, in some populations scale leaves pre-
dense, assurgent or ascending, with short ultimate dominate. Pollen cones and seed cones invariably on
branchlets, more or less glaucous. Leaves scale-like, branchlets with scale leaves. Dimensions of leaves
1.53 0.71mm, appressed, obtuse, or longest and cones not different from var. sabina.
(23mm) leaves with free apex, acute to pungent.
Seed cones more or less broad pyriform, 58mm, Taxonomic notes
with (1)24(5) seeds, yellowish glaucous.
The reduction to a variety of J. davurica Pall. under
Distribution the widespread species J. sabina L. is discussed and
explained in the Monograph of Cupressaceae and
China: N Gansu, Nei Mongol [Inner Mongolia], Sciadopitys (Farjon, 2005a: 369).
Qinghai, Shaanxi; Mongolia.
TDWG codes: 36 CHI-NM CHN-GS CHN-SX CHQ Distribution
37 MON
China, Heilongjiang, Nei Mongol [Inner Mongolia];
Ecology North Korea; Russia, E Siberia, Russian Far East;
Austria and probably occasionally elsewhere within
Artemisia steppes and grass steppes, sand dunes, the wide range of the species.
sparse vegetation on flood plains of scree and gravel, TDWG codes: 11 AUT-AU 30 (?) 31 AMU KHA PRM
S-facing mountain slopes, canyons and escarpments, 36 CHI-NM CHM-HJ 38 KOR-NK
forming patches of dwarfed scrub with herbaceous
plants and grasses. The altitudinal range is 21503350 Ecology
m a.s.l.
Juniperus sabina var. davurica is a decumbent or low
Conservation shrub in more or less open conifer forests with Larix
gmelinii, Picea obovata or P. koraiensis. Outside these
IUCN: LC forests it occurs along rocky streams as well as on
open slopes, sometimes between stands of Pinus
pumila, more often in alpine meadows and among
rocks. Its main stem is usually buried in (peaty) top
soil and in exposed situations it is extremely decum-
bent. The altitudinal range is 4001400 m a.s.l. Rock
types vary from granitic, metamorphic or volcanic
to calcareous; soils are usually peaty or leached and
acidic. The climate is maritime temperate near the
coast and in N Korea, but becomes increasingly
figure 121.Falcati
folium falciforme trees in
the Crocker Range,
Borneo
figure 123.Falcati
folium falciforme seedling
at Frasers Hill, Malaysia
figure 124.Falcati
folium taxoides on Mt.
Pani, New Caledonia
figure 125.Fitzroya
cupressoides in the N.P.
Alerce Andino (photo
P. Woltz)
figure 128. Glyptostrobus pensilis pollen
cones and seed cones (photo D. White)
figure 131. Halocarpus biformis foliage figure 132. Juniperus californica in Anza Borrego Desert State Park,
California
figure 133.Juni
perus californica seed
cones
figure 134.Juni
perus chinensis var.
sargentii foliage and
seed cones
figure 135. Juniperus communis var. figure 138. Juniperus deppeana var. figure 141. Juniperus oxycedrus subsp.
communis foliage and seed cones deppeana trunk with bark macrocarpa foliage and seed cones
figure 140.Juni
perus occidentalis var.
australis tree in the
Sierra Nevada
figure 137.Juni
perus deppeana var.
deppeana in Puebla,
Mexico
figure 142. Juniperus phoenicea subsp. phoeni- figure 143. Juniperus phoenicea subsp.
cea at Cape St. Vincent, Portugal phoenicea foliage and cones
figure 147.Juniperus
semiglobosa in Kirgyzstan
figure 151.Keteleeria
davidiana var. davidiana
foliage and seed cones
figure 150.Juniperus
virginiana var. virginiana
tree in North Carolina, USA
figure 152.Keteleeria
davidiana var. davidiana
seed cone
figure 161.Larix
lyallii in the Wenatchee
Mts., Washington, USA
figure 174.Nageia
wallichiana leaves and
seed cones (photo
L. Averyanov)
figure 175.Neo
callitropsis pancheri figure 176.Neocallitropsis
in New Caledonia pancheri foliage
continental further west, with long and cold winters ally near base and scattered stomata adaxially from
and deep snow especially at higher altitudes. This base to apex; glands present on older leaves and most
variety has also been found in a few localities in the leaves of ultimate branchlets, not active, central or
Alps, where conditions are similar to the localities in near leaf base, oval and flat; leaf colour greyish green,
mountains away from the coast in the Far East of the sometimes yellowish green. Pollen cones terminal,
species range. solitary, ovoid-oblong, 23.5mm long; microsporo-
phylls 810(12), decussate, peltate; margins minutely
Conservation denticulate, keeled towards the obtuse apex, bear-
ing 34 abaxial pollen sacs. Seed cones terminal on
IUCN: LC very short branchlets, maturing in one year, when
full grown globose or ovoid-globose, 57 4.57(8) 473
mm, sometimes wider than long and slightly bilobed,
Juniperus saltillensis M. T. Hall, Fieldiania Bot. wine-red with thick whitish blue bloom (pruinose).
34 (4): 45. 1971. Juniperus ashei J. T. Buchholz var. Bract-scale complexes in 2 decussate pairs (total 4),
saltillensis (M. T. Hall) Silba, Phytologia Mem. 7: rarely including an upper whorl of 3 (total 5), com-
32. 1984. Type: Mexico: Coahuila, Sierra Madre pletely fused, with minutely exserted triangular
Oriental, ca. 30 km SE of Saltillo, M. T. Hall 66305 bract tips; surface smooth (rugose in sicco); fibrous
(holotype F). and resinous inside. Seeds 12(3) per cone, (broad)
ovoid or ovoid-oblong, 45(6) 33.5mm, grooved
Etymology or 2-ridged, with resin pits towards the base; hilum
a third to half of length of seed, lighter than (dark)
The species epithet refers to Saltillo, a town in brown seed coat.
Coahuila, Mexico, from the vicinity of which the
species was first described. Distribution
Distribution
Ecology Distribution
Juniperus virginiana var. virginiana is a very wide- SE USA: Alabama, Florida, Georgia, Louisiana,
spread conifer in the eastern United States, extend- Mississippi, North Carolina, South Carolina (near
486 ing both its area of extent (total range) and its area the coast).
of occupancy. This happens primarily through colo- TDWG codes: 78 ALA FLA GEO LOU MSI NCA SCA
nisation on disturbed ground which is subsequently
left to settle; foremost are abandoned arable fields Ecology
and pastures, second in importance are verges of
highways, old mine tailings, etc. Other tree species Juniperus virginiana var. silicicola is restricted to a
invading these fields are Pinus spp., Ulmus ameri- narrow coastal strip near sea level of the Atlantic
cana, Populus grandidentata, Sassafras albidum, and Ocean and the Mexican Gulf and the lowlands of
Zanthophyllum americanum. In the primary range of central Florida. It is there associated with limestone
this more continental and upland variety, the most and prehistoric shell middens, often bordering tidal
commonly associated trees are Pinus virginiana, P. marshes or occurring in old dunes. Pinus elliottii,
echinata, Quercus alba, Q. rubra, Carya spp., and Quercus virginiana, Q. laurifolia, Magnolia grandi-
Juglans nigra, all of these frequently on shallow, rocky flora, Persea borbonia, and Ilex opaca are commonly
or sandy soils (glades). It is also common in moister associated trees. Over large areas the small palm
places near streams, especially in the western part of Sabal palmetto can dominate the understorey, but a
its range bordering the Great Plains, and can grow more diverse shrub layer occurs especially on more
well on a range from acid to neutral soils. Annual stabilised sites. Past selective cutting of this variety
precipitation is less than for var. silicicola, with an (Southern Red-cedar) has led to dominance of oaks
effective summer drought limit of around 350mm. and pines in many former cedar forests. Further
inland, var. silicicola becomes a minor component of
Conservation pine forests and in some areas it is sympatric with
the more widespread var. virginiana. The climate is
IUCN: LC subhumid to humid (10001600mm of rain p.a.)
with very mild winters.
Southern red-cedar, Red-cedar, Sand cedar, Coastal W. C. Cheng (Wanjun Zheng) described Juniperus
red-cedar, Coast juniper gaussenii when he was a research fellow at the
Laboratoire Forestier de Toulouse, France working uncommon crown shape not found in any true spe-
under H. Gaussen. Its type, F. Ducloux 3928 (holo. P) cies. Verification of herbarium collections in KUN
is one of several gatherings Ducloux made in and PE confirmed the cultivated status of J. gaussenii
Kunming (Yunnan-sen) early in the 20th century. in all cases where data on habitat were given. Serious
Cheng only cites one further collection, R. P. Maire doubts therefore arise concerning the status of this
s.n., coll. 10 Aug 1921, which had pollen cones and species, which is probably best regarded as a cultivar
fruit roux so perhaps indicating a monoecious of possible hybrid origin.
plant (if gathered from a single tree). On this basis
Cheng (op. cit.) compared his new species with J. Distribution
chinensis, to which it is in his opinion very close.
Differences cited are very slight though, and some, China: Yunnan, only known from cultivation in 487
e.g. the colour of seed cones, an artifact resulting some numbers in and around Kunming and in
from comparison of very few specimens. Cheng had Xichou in SE Yunnan. Also present in other ancient
no experience with living plants of his species; had cultural centres in Yunnan, e.g. Dali.
he seen them he might have concluded that there Ecology: Not known to occur in the wild and
was much less affinity with J. chinensis. Numerous apparently producing very few seed cones, most of
plants in the Botanic Garden of the Institute of which may be sterile.
Botany, Chinese Academy of Sciences, Kunming (it
is even used in hedges there), labeled as J. gaussenii, Uses
show characteristics in growth and habit, including
foliage, suggesting affinity with J. squamata as well A highly ornamental shrub which can attain
as with J. chinensis. These plants strongly suggest large size; it is not known in cultivation outside
a cultivar, which fits well with the virtual absence China.
of cones (retained juvenile stage of leaves) and an
Keteleeria Carrire, Rev. Hort. 37: 449. 1866. Type: Keteleeria fortunei (A. Murray
bis) Carrire (Pinaceae).
Yunnan yushan (Chinese); Du sam ni dt, Ngo China: Hainan Island, SW Sichuan, Yunnan; Lao
tng (Vietnamese) PDR; Viet Nam.
TDWG codes: 36 CHC-SC CHC-YN CHH 41 LAO
Description VIE 491
Description Ecology
Trees to 30 m tall, d.b.h. to 11.5 m; trunk mono- Keteleeria fortunei occurs in the hills or low moun-
podial, straight, often short and branching low; tains of SE China, in the red and yellow earth region
bark becoming thick and fissured, dark grey brown. (Wang, 1961), at elevations between 380 and 1200 m
Branches of first order heavy, long, spreading wide; a.s.l. The climate is humid, warm temperate to sub-
crown broad, often dome shaped. Branchlets slen- tropical, with annual precipitation between 1300
der, firm, (light) reddish brown or yellowish brown, and 2000 mm. It occurs in two forest formations:
glabrous, or rarely with some short hairs in grooves; the mixed mesophytic forest, and, more usually, the
leaf scars small, circular. Vegetative buds ovoid coni- evergreen broad-leaved forest. Besides many angio-
cal or subglobose, 35 24 mm, not resinous; bud sperm trees, such as evergreen sclerophyllous oaks
scales triangular, obtuse and appressed, persisting and lauraceous trees, a few additional gymnosperms
several years. Leaves spreading at 4590 from shoot, are also found in the latter formation: Pseudotsuga
(1.2)1.53(4) cm long, 24 mm wide, slightly sinensis, Cryptomeria japonica, Cephalotaxus fortu-
twisted and narrowed at base, narrowly linear to nei, and Taxus chinensis.
Conservation Uses
Although relatively widespread in distribution, The wood of this species is used locally for construc-
centuries of deforestation in southern China have tion and firewood. It is quite commonly planted in
undoubtedly reduced the forests in which this spe- China, but rare in cultivation elsewhere; it was intro-
cies naturally occurs. It is, however, capable of duced to England from seed collected in Hong Kong.
regeneration in secondary vegetation; a decline in
primary forest can therefore not be directly trans-
lated into a decline of the species.
IUCN: NT
493
494
plate 19. Keteleeria fortunei. 1. Habit of tree. 2. Branch with foliage. 3. Seed cone. 4. Immature seed cone.
5. Seed scale with seeds. 6. Seeds. 7. Leaves. 8. Leaf apices. 9. Pollen cones.
Lagarostrobos Quinn, Austral. J. Bot. 30 (3): 316. 1982. Type: Lagarostrobos franklinii
(Hook. f.) Quinn [Dacrydium franklinii Hook. f.] (Podocarpaceae).
Greek: lagaros = narrow; strobilos = cone. forward, acute. Adult leaves spirally arranged,
imbricate and appressed, rhomboid in appearance,
Description 11.5 1 mm, keeled abaxially; apex obtuse. Leaves
amphistomatic, stomata conspicuous, scattered.
See the species description. Pollen cones terminal, sessile, 46 mm long, 22.5
mm wide; microsporophylls 1015(20), rhombic to
Distribution triangular, with minutely denticulate upper margins 495
and with two basal pollen sacs containing trisac-
As for the species. cate pollen. Seed cones terminal on decurved short
branchlets, 45 mm long, consisting of 58(10)
spirally arranged fertile bracts, each with a single
Lagarostrobos franklinii (Hook. f.) Quinn, Austral. erect ovule on the adaxial side. Seeds up to 58 per
J. Bot. 30 (3): 316. 1982. Dacrydium franklinii Hook. cone, usually fewer, morphologically erect but topo-
f., London J. Bot. 4: 152, t. 6. 1845. Type: Australia: graphically pendent, ca. 2.2 2 mm, dorsiventrally
Tasmania, Huon River, [grows at Mcq Harb], A. compressed to nearly rounded in cross-section,
Cunningham s.n. (a-c) (lectotype K). Fig. 153, 154 notched ar apex, light brown, enclosed at base in a
dry, papery epimatium.
Etymology
Distribution
The species epithet commemorates Sir John
Franklin, a Governor of Tasmania in the early years Australia: Tasmania (mainly S & W parts, along
of the colony. rivers).
TDWG codes: 50 TAS
Vernacular names
Ecology
Huon pine
Lagarostrobos franklinii is mostly a riparian spe-
Description cies, usually on river banks or close to rivers, but
occasionally occurring on wet hill sides away from
Evergreen, predominantly dioecious trees to 25 m or main water courses in temperate rainforest. It forms
perhaps 30 m tall; trunk to 1.5(2) m d.b.h. (few trees groves dominated by Huon pine, marking stream
of this size now exist). Bark becoming longitudinally courses at low altitudes from sea level to 150 m, and
fissured, fibrous, exfoliating in scales and strips, ulti- it grows on some hills to 750 m in the mountain-
mately 46 cm thick, grey-brown. Crown of young ous west country. In a few areas it is observed to
trees more or less conical or pyramidal, of old mature have spread by layering, most notably on Mt. Read,
trees spreading, with large ascending main branches. where several hectares are believed to be occupied
Frequently layering, sometimes extending over large by stems forming a single clone. It is often accompa-
areas. Foliage branchlets slender, 11.2 mm diam. nied by Nothofagus cunninghamii, Eucryphia lucida,
including scale leaves, long on seedlings and young Anopterus glandulosus and ferns, with Eucalyptus
plants, or on shaded, pendulous branches, short and obliqua growing nearby on higher ground.
more rigidly spreading on sun-exposed branches in
the crown of mature trees. Juvenile leaves on seed- Conservation
lings and young plants, spirally arranged, decurrent,
spreading to all sides with free apex, 12 mm long, This species is not threatened under current (2001)
keeled abaxially, concave adaxially; apex curved IUCN criteria; it was listed as LRcd in the Conifer
Action Plan (Farjon & Page, 1999), but the subcat- Uses
egory conservation dependent is no longer recog-
nized. An estimated 15% of its habitat has been lost Huon pine was once the most important timber tree
through inundation for hydroelectric schemes and of Tasmania, but its exploitation was unsustainable
to fire over the past 100 years or so. Extensive log- and resources of good timber trees were exhausted.
ging in the past has removed nearly all large trees, The timber was exported in the colonial period of
but there is regrowth nearly everywhere; ca. 85% of the 19th century and convicts were employed in the
the remaining area of occupancy (AOO) of Huon logging and transport operations by river. Its wood
pine is now protected in reserves. One stand of the is hard and durable and was mainly used for boat
species has been made available for access to craft building and decks of sailing ships. Today, virtually
496 wood from dead and downed timber, but there is no all large trees have gone, but a few escaped the log-
cutting of living Huon pine allowed. Fire manage- gers and are now assidiously protected. A limited
ment appears to be the main priority at present to amount of down and dead wood is now used for
ensure its continued conservation under the cur- wood crafts and cabinet work. The species is rare in
rently prevailing policies. cultivation, but present in a few botanical collections
IUCN: LC and arboreta. This tree grows very slow, both in the
wild and in cultivation, and is tolerant of light frost.
Larix Mill., Gard. Dict., Abridg. Ed. 4, vol. 1. 1754. Type: Larix decidua Mill.
(Pinaceae).
Larix is the classical Latin name for larches. Section Larix with species L. decidua (type), L.
sibirica, L. gmelinii, L. czekanowskii, L. laric-
Description ina and L. kaempferi
Section Multiserialis Patschke with species L.
Monoecious, deciduous trees with a monopodial griffithii (lectotype), L. potaninii, L. lyallii, L.
trunk. Resin canals in wood, leaves and seed cones. occidentalis and L. mastersiana
Branches at regular intervals on trunk, spreading and 497
assurging or highest order branches drooping to pen- This classification was adopted im my book Pinaceae
dulous (Rauhs model). Bark thick, scaly and fissured, (Farjon, 1990); but since then a number of papers
with large plates in some species. Shoot dimorphism have been published attempting to reconstruct
pronounced; lateral short shoots give rise to leaves, a phylogeny of the genus. One of these, based on
long shoots and reproductive organs. Leaves nar- morphological data and including evidence from
rowly linear, (sub-)flexible, obtuse to acutish at apex, the fossil record (LePage & Basinger, 1995) divides
more or less flattened or broad triangular, sometimes the genus in two groups: one with short bracts and
diamond shaped in cross section, hypo- or amphis- one with long bracts. This is a similar classification
tomatic. Pollen cones solitary at apex of short shoots as the one above, but with L. kaempferi transferred
and often numerous; microsporophylls with 2 pollen to section Multiserialis; the fossils are all placed
sacs containing globular, smooth pollen with a nar- with the other, short-bracted group. The results of
row equatorial ridge. Seed cones solitary at apex of two analyses of molecular (DNA sequence) data
short shoots on a curved peduncle, more or less erect, appear to give results that group species accord-
persistent, falling attached to branches. Bracts short ing to their geographical distribution when based
and hidden or long and exserted. Seed scales rounded on plastid DNA (Wei & Wang, 2003) and nuclear
or emarginate, more or less convex, with a short pedi- ribosomal (ITS) DNA (Wei & Wang, 2004). Here,
cellate base and persistent. Seeds ovoid, held in a shal- there is a North American group (L. occidentalis,
low cup covering one side of the seed, which extends L. laricina), a North Eurasian group and a Sino-
in a relatively short, persistent wing. Seedlings with Himalayan group, but with L. sibirica behav-
57 (usually 6) cotyledons. ing aberrantly in the cpDNA based analysis and
L. lyallii (North America) not represented in the
11 species. analyses. In another study of this kind (Semerikov
et al., 2003) conflicting phylogenies were inferred
Distribution from cpDNA and ITS (with L. lyallii included).
The only consistently congruent result seems to
North America: central Alaska (disjunct); from the be a clade with the North American species L. lar-
NW Territories to Newfoundland; northern Rocky icina, L. lyallii and L. occidentalis. These belong to
Mountains and Cascade Range; New England states two different traditional sections. It appears there-
(USA). Eurasia: Europe (Alps and Carpathians); NE fore, that at present a classification informed by a
Russia across Siberia to Kamchatka and Sakhalin; robust phylogeny that would help to explain char-
Japan (disjunct), NE China; Sino-Himalayan moun- acter transformations in an evolutionary context
tain system. still eludes us. No formal classification is therefore
presented here.
Synopsis
Key to the species of Larix
The genus Larix has been divided into two groups
based on morphological characters in the seed cones 1a. Bracts of mature seed cones shorter than seed
and, to a lesser degree, in the leaves. These groups scales, usually barely visible in opened cones;
have been formalized in sections: ripe seed cones globose, ovoid or ovoid-conical.
Leaves carinate on invers-dorsal side (below) Larix czekanowskii Szafer, Kosmos 38: 1297. 1913.
only 2 Type: Russia: Siberia, Tunguska River, below mouth
1b. Bracts of mature seed cones (much) longer of Tomezoy River, A. Czekanovsky & F. Mller s.n.
than seed scales, straight or reflexed; ripe seed (lectotype LE).
cones ovoid-conical to oblong-cylindrical.
Leaves carinate on both sides, rarely only on Etymology
invers-dorsal side 7
2a. Seed scales concavo-convex, with strongly This nothospecies was named after Aleksander P.
recurved upper margin; seed cones broad Czekanowski (18331876), a Polish geologists who
ovoid, (1.5)23(3.5) cm long L. kaempferi studied the geology of the Lake Baikal Basin while
498 2b. Seed scales convex or more or less straight, with in exile.
flat upper margin; seed cones variously shaped
3 Vernacular names
3a. Seed scales 1020 per cone, not spreading; seed
cones 12 cm long L. laricina No common names have been recorded for this
3b. Seed scales (15)2040(45) per cone, spread- taxon.
ing; seed cones (1.5)1.85(6) cm long 4
4a. Leaves hypostomatic (all or nearly all stomata Description
on one side) L. gmelinii
4b. Leaves amphistomatic (stomata on both sides, Trees which are intermediate between Larix sibirica
but usually more on one side) 5 and L. gmelinii and appear to be common in a wide
5a. Seed scales of green cones (densely) pubescent, belt stretching N to NW from Lake Baikal share some
becoming more glabrous with age L. sibirica character states of both species, particularly evident
5b. Seed scales of green cones always glabrous 6 in the cones. The seed cones are smaller than those
6a. Seed scales incurved, straight or slightly of L. sibirica and resemble those of a large-cone vari-
recurved; upper margin repand or emarginate, ety of L. gmelinii: var. principis-rupprechtii; the seed
sometimes entire (European species) scales are less pubescent than those of L. sibirica.
L. decidua They have been interpreted by Russian botanists and
6b. Seed scales incurved; upper margin entire foresters as of natural hybrid origin.
(Siberian species) L. czekanowskii
7a. Apex of bracts in seed cones broadly acute Distribution
L. potaninii
7b. Apex of bracts in seed cones cuspidate, often Central Siberia: from Lake Baikal to the mouth of
narrow and elongated 8 the Yenisei River
8a. Mature seed cones 58(11) cm long, cylindrical TDWG codes: 30 irk kra
L. griffithii
8b. Mature seed cones 2.55(6) cm long, variously Ecology
shaped but longer than wide 9
9a. Leaves amphistomatic. Bracts straight, with an Larix czekanowskii occurs in central Siberia, where
abruptly narrowing cusp L. lyallii it forms taiga forest with Picea obovata, Pinus sylves-
9b. Leaves hypostomatic. Bracts recurved (at least tris and broad-leaved trees such as Betula pendula
the cusp) 10 and Populus spp., broadly following the Yenissei
10a. Seed scales recurved, mostly emarginate (some- River. It grows on a great variety of soils, from peat
times entire); bracts with a thin, elongated bogs to well drained, sandy or rocky soils, where
cusp L. occidentalis it has its optimum. The climate is very cold (min.
10b. Seed scales more or less straight, entire or temp. 55 C), continental or subarctic, dry, with
slightly emarginate; bracts short cuspidate very long winters.
L. mastersiana
Conservation mata on lower side in two narrow bands separated
by a keel; leaf colour light green, darkening, yellow
IUCN: NE in autumn. Pollen cones terminal on short shoots,
numerous on pendulous long twigs, 0.51 cm long,
Uses yellow, perular scales with fimbriate margins, red-
dish. Seed cones terminal on short shoots, turning
The wood of this larch is durable and used in con- erect; peduncles curved, 0.51 cm long, subtended
struction, traditionally for log houses in Siberia by leaves; cones ovoid or ovoid oblong, with obtuse
for which the wood is roughly hewn to shape, but apex, (1.2)2.54(4.5) cm long, (1.2)1.53(3.5) cm
untreated. It has been widely used for railroad sleep- wide with opened scales; colour (immature) dark
ers e.g. on the famous Trans Siberian Railroad. Larch red or purplish, sometimes green, maturing to pale 499
wood is also milled for construction timber and green with purplish margins of seed scales, ripen-
veneer, and pulped for the paper industry. This natu- ing to (dark) brown, old cones grey. Seed scales
ral hybrid larch is not known to be in cultivation, but 2535, ovate to suborbicular, slightly convex, 715
it may be present in e.g. Scandinavia under the name 613 mm; surface striated, shining smooth in older
L. sibirica. cones, reddish pubescent near base, later glabrous;
upper margin entire, incurved, repand or emargin-
Larix decidua Mill., Gard. Dict., ed. 8: Larix No. 1. ate, sometimes slightly recurved; base narrowed.
1768. Bracts ligulate, with acicular or cuspidate apex,
length - seed scales, mostly included, but vis-
Etymology ible with opened scales. Seeds ovoid-cuneate, 4 2.5
mm, dark brown-grey; seed wings oval, 610 46
The species epithet refers to the deciduous (season- mm, light brown.
ally falling) leaves.
Distribution
Vernacular names
Europe: Alps, Carpathians, Slovakian Mts., S Poland.
European larch; Gemeine Lrche (German); Mlze TDWG codes: 11 AUT-AU AUT-LI CZE-CZ CZE-SL
dEurope (French) GER POL SWI 12 FRA-FR 13 ITA-IT ROM YUG-SL 14
UKR-MO UKR-UK
Description
Ecology
Trees to 4050(55) m tall, d.b.h. to 1.52.5 m; trunk
monopodial, straight or curved at base; bark deeply Larix decidua occurs in the high mountains of cen-
fissured, breaking into large plates, flaking, expos- tral Europe, at altitudes between (600)10002200(
ing reddish inner bark. Branches heavy, long, curved 2500) m a.s.l., in the Central Alps it usually forms the
down, ascending near the ends; branches of second tree limit. The soils are neutral to acidic, mostly on
order long, slender, pendulous; crown pyramidal in granitic rock. The climate has cool, moist summers
young trees, widening in old trees. Branchlets thin, and cold, snowy winters, but annual precipitation
slender, flexible, or stout, pink buff or (pale) yel- rarely exceeds 1000 mm. Pure stands are uncom-
lowish; glabrous, or slightly pubescent when young; mon, more often it is mixed with Pinus cembra in
short shoots cylindrical or subglobose, 0.31 cm the Alps, below 1800 m also with Picea abies.
long. Vegetative buds ovoid, 3 2 mm, not resin-
ous. Leaves on long shoots remote, appressed for- Uses
ward against the shoot or spreading; on short shoots
crowded in false whorls, 3040, (2.33.5(4) cm long, The wood of European larch is valued for its durabil-
0.51 mm wide, narrowly linear, soft, flexible, in ity and has been used for centuries in the Alps and
cross-section broad triangular or flattened, carinate Carpathians to build houses; traditional style houses
below, obtuse or acutish at apex; amphistomatic, sto- still use well seasoned wood of this species. Other
traditional uses are fences, gates, feeding racks, and Distribution
water troughs for animals. Due to its durability the
wood of European larch has been used extensively Central to East Europe: Sudeten, Tatra, Carpathians.
for railway sleepers, until these were replaced by TDWG codes: 11 CZE-CZ CZE-SL POL 13 ROM 14
concrete and iron structures in modern times. Trees UKR-MO UKR-UK
with a curved base were split and hollowed and the
two halves joined to make Alphorns, large wind Conservation
instruments with a far carrying low tone; competi-
tions to blow the horn are still held in some regions IUCN: LC
of the Alps. This species has been introduced in the
500 lowlands of Europe for plantation forestry as well as Larix decidua Mill. var. polonica (Racib. ex
an amenity tree. In horticulture for gardens it is not Wycicki) Ostenf. & Syrach, Pflanzenareale 2: 63.
so common, although a modest number of cultivars 1930. Larix polonica Racib. ex Wycicki, Obraz.
is known, most with various branching habits. Rosl. Krl. Polsk. 2: 1516, t. 1. 1912. Type not
designated.
3 varieties are recognized:
Description
Larix decidua Mill. var. decidua. Type: Illustration
Larix folio deciduo Conifera in Miller, Cat. Pl.: 43, Long shoots pale yellowish; leaves flattened. Seed
t. 11. 1730 (lectotype). Fig. 155, 156, 157 cones ca. 1.5 1.2 cm, with suborbicular, convex
scales with a round, entire upper margin.
Description
Distribution
Seed cones 2.54.5 cm long, 1.53 cm wide when
opened; seed scales with a rounded or more or less Poland (headwaters of Wista River).
repand to emarginate upper margin. TDWG codes: 11 POL
Distribution Ecology
Europe: Alps, Carpathians, Slovakian Mts. This variety has been found in scattered stands in
TDWG codes: 11 AUT-AU AUT-LI CZE-CZ CZE-SL mixed forests at ca. 150350 m a.s.l. It is commonly
GER SWI 12 FRA-FR 13 ITA-IT YUG-SL associated with Pinus sylvestris and Betula pendula,
sometimes with Quercus robur.
Conservation
Conservation
IUCN: LC
In forests surrounding the headwaters of the Wista
Larix decidua Mill. var. carpatica Domin, Sborn. River in Poland, larches have become very rare. In
Vyzk. Ustav Zemed. R..S. 65: 149. 1930. Type: most areas only ancient relict trees survive and there
Slovakia, Tatra Mts., V. Krajina s.n., 7 sep 1931 is little or no successful regeneration. This could
(holotype PR). partly be the result of historical climate change,
helped by forest management practice and land uses
Description that have favoured pioneer broad-leaved trees and
pines. The geographical extent of this variety is not
Seed cones (1.2)22.2(2.5) cm long, about as wide exactly known but its area of occupancy (AOO) is
when opened; seed scales often with an entire, less than 500 km.
rounded upper margin. IUCN: EN [B2ab (ii, iv, v)]
Larix gmelinii (Rupr.) Kuzen., Trudy Bot. Muz. reddish pubescent, usually weakly pubescent or gla-
Rissijsk. Akad. Nauk 18: 41. 1920. brous; upper margin entire, rounded or truncate,
or weakly to strongly emarginate and slightly or
Etymology strongly recurved; base narrowed or slightly pedi-
cellate. Bracts broad ligulate-lanceolate, length
This species was named after Johann Georg Gmelin seed scales, included except at base of cone, but
(17091755), a botanist who traveled widely in visible with opened seed scales; apex trilobate with
Siberia. longer cusp. Seeds ovoid, slightly flattened, 23 12
mm, light brown; seed wings ovate-oblong, 56 3
Vernacular names mm, bright orange-brown, shining.
501
Dahurian larch; Listvennitsa daurskaya (Russian); Distribution
luoye song (Chinese); gui-matsu (Japanese)
NE China: Hebei, Manchuria, Nei Monggol [Inner
Description Mongolia], N Shanxi; North Korea; NE Mongolia;
Russian Federation: E Siberia, Russian Far East;
Trees to 3035 m tall, d.b.h. to 11.5 m; trunk mono- Japan (Hokkaido).
podial; bark on trunk dark red brown, with greyish TDWG codes: 30 BRY CTA IRK YAK 31 AMU KAM
plates, finely scaly. Branches long, spreading hori- KHA KUR MAG PRM SAK 36 CHI-NM CHM CHN-HB
zontally, ascending near the top; branches of second CHN-SX 37 MON 38 JAP-HK KOR-NK
order relatively short, firm, drooping, not pendulous;
crown broad pyramidal, open, flat topped and irreg- Ecology
ular in old trees; Branchlets thin, slender, light pink
brown to orange-brown or dark purplish brown; Larix gmelinii occupies a very large area and there-
pubescence variable, weak or dense, or glabrous; fore occurs in a wide range of habitats: on lowland
short shoots small, 36 mm, cylindrical. Vegetative subarctic plains, in river valleys, in mountains and
terminal buds ovoid globose, with acute apex, not also on the edges of moors and swamps. Its altitudi-
or slightly resinous; bud scales ovate or obtuse tri- nal range is between 300 m and 1800 m a.s.l. The soils
angular, dark brown or purplish brown. Leaves on are affected by permafrost over much of the range;
short shoots crowded in false whorls, 2030(35), the climate in large parts of its range is continental
1.53(4) cm long, 0.50.8 mm wide, narrowly lin- subarctic, with very cold winters, and relatively dry
ear, widest near apex, soft and flexible, cross-section (400 mm to 500 mm in the Greater Hinggan Range)
diamond shaped, or flattened and keeled below; to extremely dry in the subarctic interior. In more
apex obtuse or acutish; stomata in two narrow bands maritime areas of the Russian Far East precipitation
separated by a keel below, none or a few above; leaf rises to 1000 mm and more. It is the only tree species
colour bright green, yellow in autumn. Pollen cones in E Siberia reaching the tree limit at 70 N, but in
terminal on short shoots, 57 mm long, yellow. Seed the more southern part of its range it is commonly
cones terminal on short shoots, turned erect; pedun- mixed with Abies sibirica, Picea obovata and Pinus
cles short, curved; cones ovoid or ovoid-oblong, sylvestris. In the boreal coniferous forest it is a cli-
widening with opened scales; apex truncate, 0.8 max species only on permafrost or peaty soils. In the
3.5(4.5) cm long, 0.82(3) cm wide with opened high swamps and bogs in Sakhalin and on the Kuril
scales; colour (immature) purplish red or sometimes Islands it usually forms pure stands, but on some-
light green, maturing to rose-purple or greenish pur- what drier sites it is mixed with Abies sachalinen-
ple, ripening to bright or dark red brown; old cones sis var. sachalinensis, Picea jezoensis, Alnus hirsuta,
grey. Seed scales (15)2040(45), ovate oblong, Betula japonica, B. ermanii, and Salix spp. In central
slightly recurved or straight, convex, flattening out Kamchatka occurs a relict taiga forest with a mixture
when opened, 510 48 mm at mid-cone; surface of Larix gmelinii and Picea jezoensis, accompanied
smooth, striated on abaxial side, sometimes strongly by Betula sp. and with Pinus pumila and Juniperus
communis var. saxatilis common in the understo- brous or weakly pubescent; upper margin from entire
rey. In mountain ranges on the maritime seaboard and rounded to emarginate and slightly recurved.
of NE Asia conditions for mixed coniferous forest
are more optimal and here L. gmelinii grows with Distribution
Abies nephrolepis, A. holophylla, Picea obovata or
P. jezoensis, and Betula ermanii, locally also with NE. China: Manchuria, Nei Monggol [Inner
Juniperus, Pinus pumila or Rhododendron sp. in the Mongolia]; Mongolia; Russian Fed.: E. Siberia,
understorey. Russian Far East (including Kamchatka).
TDWG codes: 30 BRY CTA IRK YAK 31 36 CHI-NM
Uses CHM 37 MON
502
Dahurian larch is an extremely important timber Conservation
tree in the Russian Far East, where it provides build-
ing logs for traditional log houses, railway sleepers, IUCN: LC
fences, and gates, as well as timber for construction,
ship building (in Japan and Sachalin), and the pulp
industry. Its variability also extends to the suitabil- Larix gmelinii (Rupr.) Kuzen. var. japonica
ity as a plantation tree for forestry outside its natural (Maxim. ex Regel) Pilg., in Engler & Prantl, Nat.
range; provenances from Siberia are prone to late Pflanzenfam., ed. 2, 13: 327. 1926. Larix dahurica
frosts (which actually means: lack of a proper cold Turcz. ex Trautv. var. japonica Maxim. ex Regel,
winter that lasts until spring definitively arrives), Gartenfl. 20: 105. 1871; Larix gmelinii (Rupr.) Kuzen.
while those of Japan and even Sakhalin are less likely subsp. japonica (Maxim. ex Regel) E. Murray,
to be frost damaged. Inter-specific hybrids have been Kalmia 12: 21. 1982. Type: Japan: Hokkaido, Oshima
produced in cultivation in Denmark and China, but Prov., Hakodate, cultivated on temple grounds,
probably have not been planted far beyond the trial C. J. Maximowicz s.n. (holotype not located,
nurseries. Due to the risk of frost damage this spe- isotype K).
cies and its varieties are rare in cultivation in more
temperate parts of the world. Description
4 varieties are recognized: This variety differs from var. gmelinii in the follow-
ing characters: Seed cones (when opened) wider
than long, 1.22.5 1.52.8 cm; seed scales more
Larix gmelinii (Rupr.) Kuzen. var. gmelinii. Abies numerous, 1825, ovate oblong, with emarginate
gmelinii Rupr., Beitr. Pflanzenk. Russ. Reiches 2: 56. and (strongly) recurved apex. Young shoots dark
1845. Type not designated. purplish brown, densely pubescent at first. Leaves
slightly shorter: 1.52.5(3) cm.
Description Conservation
The two similar names Larix griffithii and L. griffithi- 2 varieties are recognized:
ana have been used more or less interchangeably in
the literature on conifers, in fact I used the latter in Larix griffithii Hook. f. var. griffithii. Type: Bhutan:
my book Pinaceae (Farjon, 1990). However, the for- [Bootan, locality not stated], W. Griffith 4989
mer (1854) has priority over the latter (1855), the pre- (lectotype C). Fig. 159
sumed basionym Abies griffithiana Lindley et Gord.
(1850) being a nomen nudum, and must be applied Abies griffithiana Lindl. & Gordon, J. Hort. Soc.
to this species following the International Code of London 5: 214. 1850, nom. nud.
Botanical Nomenclature. Larix griffithiana hort. ex Carrire, Trait Gn.
Conif.: 278. 1855.
Distribution Larix kongboensis R. R. Mill, Novon 9 (1): 79. 1999.
Conservation Description
Alpine larch, Subalpine larch, Lyall larch Larix lyallii is a subalpine larch which occurs at or
near tree line, at elevations between 1520 m and
Description 2440 m (max. 3020 m) a.s.l. It grows usually on shal-
low, rocky mountain soils, but occasionally on deeper,
Trees to 2025 m tall, d.b.h. to 0.50.8 m; trunk well drained soils if there are no competitive species.
monopodial; bark scaly and fissured below in old The climate is cold, with short, cool summers and
trees. Branches moderately long, slender or mas- long, snowy winters. It may occur in pure stands or
sive, ascending to nearly erect, or more horizontal; mixed with e.g. Abies lasiocarpa, Pinus albicaulis,
508 branches of second order short, flexible, not pendu- P. flexilis, Picea engelmannii, and Tsuga mertensiana,
lous; crown broad or narrow, conical, irregular and forming small groves near the tree line or scattered,
open in old trees. Branchlets rather stout, firm, orange solitary, stunted trees, sometimes surrounded by very
red, but young shoots densely covered with lanate, little vegetation taller than alpine meadows.
yellowish hairs, in second year almost glabrous, grey,
later blackish grey; short shoots ovoid-conical or Conservation
barrel-shaped, with grey, pubescent rings and apex,
510 mm long. Vegetative buds ovoid globose, 3 2 IUCN: LC
mm, not resinous, yellowish pubescent; bud scales
obtuse triangular, brown. Leaves on short shoots spi- Uses
rally, close, in false whorls of 2535(40), (1.5)23.3
cm long, 0.61 mm wide, narrowly linear, curved or Due to its usually scattered occurrence at high alti-
twisted, more or less rhombic in cross section, keeled tude Subalpine larch has no commercial value as a
on both sides, obtuse or acute at apex; amphistomatic, timber tree. Its wood properties are similar to other
but more lines of stomata below; leaf colour light larches. It establishes as a pioneer in avalanche
green, later bluish green, yellow in autumn. Pollen chutes and can minimize the destructive impact of
cones terminal on short shoots, 1015 mm long, yel- snow avalanches better than most other conifers.
low. Seed cones terminal on short shoots, more or less In horticulture, it is very susceptible to late frosts
erect; peduncles short, thick, curved; cones ovoid- and therefore seldom planted except in regions with
conical to cylindrical, obtuse or truncate at apex, 3.55 long, consistently cold winters.
cm long, 22.5 cm wide with opened scales; colour
(immature) yellowish pubescent, almost hidden by
purplish red bract scales, maturing to light or pale Larix mastersiana Rehd. & E. H. Wilson, in
brown with purplish bracts. Seed scales ovate oblong Sargent, Pl. Wilson. 2: 19. 1914. Larix griffithii Hook.
or sub-orbicular, recurved when opened, 1014 f. var. mastersiana (Rehd. & E. H. Wilson) Silba,
612 mm at mid-cone; abaxial surface densely pubes- Phytologia Mem. 7: 39. 1984. Type: China: Sichuan,
cent when young, later glabrous, finely striated or Qionglai Shan, Wolong Reserve [Kuan Hsien],
smooth; upper margin entire, rounded, not incurved; E. H. Wilson 906 (holotype A).
base cuneate or narrowed. Bracts broad ligulate-lan-
ceolate, narrowing into a long, caducous cusp, length Etymology
1.5 seed scales, exserted, straight; cusps recurved.
Seeds triangular or ovoid, flattened, 4 3 mm, light The epithet commemorates Maxwell T. Masters
brown; seed wings ovate oblong, 810 mm long, light (18331907), a physician and honorary botanist at
brown or rose-brown, tinged with red. the Royal Botanic Gardens, Kew, who contributed
much to the study of conifers.
Distribution
Vernacular names
Canada: SW Alberta, SE British Columbia; USA: N
Idaho, W Montana, N Washington. Masters larch; chuan hong shan (Chinese)
TDWG codes: 71 ABT BRC 73 IDA MNT WAS
Description survey found only the following three localities: Min
River drainage, Dadu River drainage, and upstream
Trees to 2025 m tall, d.b.h. to 0.8 m; trunk mono- of Qingyijiang (State Forestry Bureau, 2009).
podial, straight or curved; bark on trunk irregularly TDWG codes: 36 CHC-SC
fissured and breaking into grey plates. Branches
long, the topmost ascending, the lower spread- Ecology
ing horizontally or descending; branches of second
order slender, pendulous, but less so than those of Larix mastersiana is a high mountain species of rare
L. potaninii; crown usually broad, domed in free occurrence, its altitudinal range is between 2000 m
standing trees, or more conical in forest stands. and 3500 m a.s.l. It grows in podzolic mountain soils,
Branchlets long, slender, pendulous, yellowish or usually on steep slopes with good drainage. The cli- 509
reddish brown, becoming grey, glabrous, or very mate is cold-temperate and moist. It has not been
young shoots sparsely pubescent; short shoots cylin- reported from climax forest with other conifers, but
dric, with rings of revolute scale bases, 315 mm seems to be a tree thriving in secondary growth after
long. Vegetative buds conical or ovoid, 2 1.5 mm, forest clearing, as inferred from photographs by E.
resinous; bud scales triangular, light brown, shin- H. Wilson taken at the beginning of the 20th century
ing. Leaves on on short shoots spirally, densely set in [collection of the Arnold Arboretum, Mass., USA
false whorls of 2040, (1.2)23(3.5) cm long, 1 mm and a 4-volume photo-album entitled Vegetation
wide, narrowly linear, slightly wider near the obtuse of Western China (Wilson, 1912), copy in the
to acutish apex, more or less rhombic in cross sec- Herbarium of the Natural History Museum at Paris].
tion, keeled on both sides, most clearly near base;
stomata none or only a few faint lines above, in 2 Conservation
narrow bands below; leaf colour bright green, yellow
in autumn. Pollen cones terminal on short shoots, Exploitation beyond sustainability has led to serious
pedunculate, erect or pendant, 1015 mm long, yel- decline of this species in the more accessible parts of
low. Seed cones terminal on short shoots, more its limited natural range. It is now largely confined
or less erect from pendulous branches; peduncles to the steeper and higher localities where forest road
curved, 5 mm long; cones ovoid-cylindrical, curved building has not advanced. Recent Chinese policies
or straight; apex obtuse, 2.54.5 cm long, 1.52.5 cm to discontinue logging of the natural forests in west-
wide with opened scales; colour (immature) green- ern regions may halt or even reverse the decline.
ish, with orange yellow bracts covering seed scales, IUCN: EN [A2cd; B2ab (iiv)]
maturing to light brown, with dark brown bracts, old
cones dull, dark brown, with blackish bracts. Seed Uses
scales 3040, obcordate-orbicular, convex, 610
712 mm at mid-cone; surface smooth or slightly A timber tree used for construction, pit props, rail-
striated, puberulent when young, later glabrous; way sleepers and furniture; the bark yields tannins.
upper margin entire, slightly emarginated, base very It has been over-exploited in its natural range, espe-
short pedicellate or cuneate. Bracts broad, lanceo- cially in more accessible localities. Outside China,
late, exposed part triangulate, reflexed, with lacerate where it is used in afforestation, this species is not in
margins, cuspidate, 22.3 cm long, exserted. Seeds cultivation except for a few specimens in living tree
ovoid-cuneate, 3 2 mm, yellowish brown or light collections (arboreta).
brown; seed wings obovate, 68 45 mm, light yel-
lowish brown.
Larix occidentalis Nutt., N. Amer. Sylva 3: 143, t.
Distribution 120. 1849. Type not designated.
plate 20. Larix potaninii. 1. Habit of tree. 2. Branchlet with foliage. 3. Branch with seed cones (var.
potaninii). 4. Seed cone (var. macrocarpa). 5. Seeds. 6. Seed scale with bract.
Larix potaninii Batalin var. potaninii. Type not Larix potaninii Batalin var. himalaica (W. C. Cheng
designated. & L. K. Fu) Farjon & Silba, Phytologia 68: 37. 1990.
Larix himalaica W. C. Cheng & L. K. Fu, Acta
Description Phytotax. Sin. 13 (4): 84. 1975. Type: China, Xizang
[Tibet], north side of Xomolungma [Mt. Everest],
Seed cones 3.55.5 cm long, 1.53 cm wide when Chinese collector No. 80A (holotype PE).
scales are opened; seed scales 3565 in number.
Description
Distribution
Seed cones max. 6.5 cm long; bracts mucronate cus-
China: S Gansu, S Shaanxi, W Sichuan, NW Yunnan, pidate. Young shoots yellowish orange. 513
E Xizang [Tibet].
TDWG codes: 36 CHC-SC CHC-YN CHN-GS Taxonomic notes
CHN-SA CHT
In Flora of China 4: 34 (1999) this variety has been
Conservation treated as a distinct species. The morphological dif-
ferences appear to be of a quantitative and more or
IUCN: LC less gradual nature and it shares the typical upright
bracts of L. potaninii.
Vernacular names The correct name for this species is Larix sibirica
Ledeb. (1833) as it is the earliest binomial published
Siberian larch; Listvennitsa sibirskaya (Russian); under Larix for this species. In my book Pinaceae
xian bei luo ye song (Chinese) (Farjon, 1990) I incorrectly used L. russica (Endl.)
Sabine ex Trautv., based on Pinus larix L. var. rus-
Description sica Endl. (1847) as the accepted name. This name
was also used in Dallimore & Jacksons well known
Trees to 3040 m tall, d.b.h. to 11.5; trunk monopo- Handbook of Coniferae and Ginkgoaceae (e.g. edi-
dial, straight or curved; bark on trunk light brown, tion 4, 1966) and in other compilations of coni-
rough and scaly. Branches spreading horizontally, fers. This species was long considered a mere form
near the top ascending; branches of second order or variety of the European larch, L. decidua, first
slender, long, drooping or pendulous; crown broad described as Pinus larix by Linnaeus (1753). When
conical, often irregular. Branchlets slender, flex- raised to species rank, it is compulsory to take up the
ible, pale yellowish, lustrous, usually glabrous or earliest species epithet published for it, which was
with minute pubescence in grooves; short shoots given by Ledebour in his Flora Altaica (op. cit.).
conical-globose or short cylindrical, 410 mm long.
Vegetative buds ovoid, 3 2 mm, not resinous; bud Distribution
scales ovate, more or less pubescent, dark greyish
brown. Leaves on short shoots spirally, close, in false Russia: from the White Sea to Lake Baikal in Siberia;
whorls of 2040, (2)2.54(5) cm long, 0.51 mm China: Xinjiang (Altai Mts., E Tien Shan); Mongolia
wide, narrowly linear, soft, flexible, flattened, faintly (Altai Mts.).
keeled below, obtuse or acutish at apex; stomata TDWG codes: 14 RUN RUE 30 ALT IRK KRA TVA
in a few lines above, in two narrow bands below; WSB 36 CHX 37 MON
Ecology Uses
Larix sibirica is common in the lowland taiga of W Siberian larch is an important timber tree in Russia.
Siberia, where it forms the northern limit of trees It is logged from natural coniferous forests as well
alternately with Picea obovata and with Pinus syl- as from plantations, which are established in Russia
vestris. It also occurs in the mountains (from 500 outside its natural range as well as in Finland, Poland
m to 2400 m a.s.l.). It grows on a great variety of and Sweden. The wood is durable and used in con-
soils, from peat bogs to well drained, sandy or rocky struction, traditionally for log houses in Siberia
soils, where it has its optimum. The climate is very for which the wood is roughly hewn to shape, but
cold (min. temp. 55 C), continental or subarctic, untreated. It has been widely used for railroad sleep-
dry, with very long winters. There are pure stands on ers e.g. on the famous Trans Siberian Railroad. Larch 515
peat or on mountains above the steppe (Altai Mts.), wood is also milled for construction timber and
but more common it is mixed with Pinus sylvestris, veneer and pulped for the paper industry. Siberian
P. sibirica, Picea obovata, Abies sibirica and broad- larch is in cultivation as an amenity tree, but it is vul-
leaved trees such as Betula pendula and Populus spp. nerable to late frosts in climatic zones with mild,
fluctuating winter temperatures. For this reason it is
Conservation not used in western Europe but successfully planted
in central and eastern Europe, in Scandinavia, and as
IUCN: LC far afield as Iceland. A few cultivars are known, but
with limited distinction from habit forms that also
occur in nature.
Lepidothamnus Phil., Linnaea 30: 730. 1861. Type: Lepidothamnus fonkii Phil.
(Podocarpaceae).
Greek: lepis, lepidos = scale; thamnos = bush, shrub. 2b. Adult scale leaves on ultimate branchlets 1.53
11.5 mm, strongly keeled to gibbous. Native
Description in Chile and Argentina L. fonkii
This dwarf conifer occurs in Sphagnum bogs and Lepidothamnus intermedius (Kirk) Quinn, Austral.
moorlands; its stems are usually hidden in moss or J. Bot. 30 (3): 316. 1982. Dacrydium intermedium
other vegetation and only upright foliage branches Kirk, Trans. New Zealand Inst. 10: 386. 1878. Type:
protrude. It can form quite extensive low thickets New Zealand: South Island, Westland, Hokitika, T.
that exclude all or most other plants, rooting on the Kirk 806 (syntype K, lectotype not designated).
decumbent stems. Its altitudinal range varies with
latitude as it occurs roughly between 40 S and 55 S, Dacrydium intermedium Kirk var. gracilis Kirk,
between 900 m in the north to near sea level in the Trans. Proc. New Zealand Inst. 17: 224. 1885. Type:
south. In the north it is growing in bogs in forested New Zealand: Stewart Island, Ruggedy, T. Kirk 1072
areas with Pilgerodendron uviferum, Fitzroya cupres- (lectotype K, designated here).
soides and Nothofagus antarctica. The latter antarctic
beech is virtually the only tree at its southern limit, Etymology
in between Pilgerodendron uviferum also occurs fre-
quently. Sedges (Carex spp.) and other plants char- The species epithet refers to intermediacy of char-
acteristic of these bogs are also common. acters compared to other species of Lepidothamnus.
Ecology Description
plate 21. Lepidothamnus laxifolius. 1. Habit of shrub. 2. Branch with foliage and seed cones. 3. Branchlet
with juvenile leaves. 4. Juvenile leaves. 5. Adult leaves. 6. Pollen cone. 7. Seed cone and seed.
1 m long, occasionally branching. Foliage branches Ecology
very slender and flexuous, 12 mm thick, spread-
ing or ascending, sparse or more crowded. Leaves of Lepidothamnus laxifolius is one of the smallest coni-
two types, juvenile and adult (intermediates occur), fers. It occurs in moorlands, peat bogs, and tus-
often mixed on a single leading branch but adult sock grass slopes in the mountains, usually between
leaves distal from juvenile leaves. Juvenile leaves 750 m and 1200 m a.s.l., on Stewart Island down to
alternate, subulate to linear, stiff and spreading at sea level. It can form extensive mats creeping over
more or less right angles to shoot, 38 mm long, rocks and long runners trailing through the vegeta-
keeled abaxially, flat or slightly concave adaxially, tion. It is associated with tussock grass (Chionochloa
obtuse. Transitional leaves shorter but similar, 1.53 rubra) and with Halocarpus bidwillii, H. biformis,
520 mm long, spreading at less than 90. Adult leaves Podocarpus nivalis, Hebe (Veronica s. l.), Olearia,
spirally arranged, imbricate, appressed, oblong- Dracophyllum, Pseudopanax, Gleichenia, Phormium,
ovate, 11.5 1 mm, slightly keeled, obtuse. Stomata and other shrubby plants and ferns.
on juvenile leaves numerous on adaxial side, in two
sparse bands on abaxial (keeled) side; scattered on Conservation
adult leaves. Pollen cones solitary, terminal, sessile,
57 2 mm; microsporophylls broadly triangular IUCN: LC
with acute-apiculate apex and two basal red pollen
sacs. Seed cones solitary, terminal, consisting of a Uses
short axis with 3 slightly spreading bracts that fuse
and swell to a globose, succulent (but sometimes dry Pygmy pine has no commercial value, but would be
and not swollen) receptacle and turn orange-red. an interesting and probably excellent dwarf shrub
Seeds enclosed at base by an epimatium, 45 mm for rock gardens. It is currently only present in a few
long, ovoid-oblong, with a small, curved apiculus, botanic gardens and perhaps an occasional private
ripening brown or purplish brown with longitudinal collection. With plants only 78 cm long known to
lighter striations. have borne ripe seeds, it may well be the smallest
conifer species known.
Distribution
Stegocedrus Doweld, Novosti Sist. Vyssh. Rast. 33: 42. sequences (Gadek & Quinn, 1993; Brunsfeld et al.,
2001. Type: Stegocedrus austrocaledonica (Brongn. & 1994) and on matK sequences (Gadek et al., 2000) it
Gris) Doweld [Libocedrus austrocaledonica Brongn. & was found that there is no close relationship between
Gris]. Libocedrus s. str. and Calocedrus, the northern hemi-
sphere genus that was long included. Phylogenetic
Greek: libos = tear, drop (referring to resin exuda- analysis based on morphology as well as on com-
tions); Cedrus is the classical name for (true) cedars. bined data (Gadek et al., 2000) confirmed this, so 521
there is strong support for at least the separation
Description of these two. Phylogenetic relationships are much
closer with the southern hemisphere taxa, but how
Shrubs or trees to 35 m, evergreen, monoecious, mul- close and with which taxa has not been unambigu-
tistemmed or monopodial. Bark scaly, exfoliating in ously demonstrated. Here the data have so far pro-
longitudinal strips or plates, reddish brown or brown. duced conflicting results in phylogenetic analysis.
Branches spreading or ascending, forming a pyrami- Molecular data have placed Pilgerodendron nested
dal, conical or bushy crown. Foliage branches fron- within Libocedrus (Gadek et al., 2000) but morphol-
dose and plagiotropic, sometimes more irregularly ogy did not (Hill & Brodribb, 1999; Farjon, 2005a).
disposed, spreading or ascending. Leaves scale-like,
decussate, decurrent, imbricate, strongly dimorphic Key to the species of Libocedrus
on flattened (pen)ultimate branchlets, facials smaller
than laterals or nearly equal in size; facials rhombic, Leaves should be examined on ultimate branchlets
15 mm long; laterals divergent, 27 mm long, condu- of full grown, not young plants.
plicate, falcate, both eglandular; margins entire; apex
free or appressed, obtuse to acute; stomata predomi- 1a. Large trees. Pollen cones with 814 microspo-
nantly in conspicuous bands on underside of branch- rophylls. Native to New Zealand 2
lets. Pollen cones terminal, solitary, 2.510 23.5 mm; 1b. Shrubs or small trees. Pollen cones with 1624
microsporophylls 824, decussate, peltate, bearing microsporophylls. Native to New Caledonia 3
4(6) abaxial pollen sacs. Seed cones terminal on flat- 2a. Ultimate branchlets more or less quadrangular,
tened or quadrangular branchlets, subtended by 45 with facial leaves only slightly smaller than lat-
transitional leaf pairs, 818 mm long; bract-scale com- eral leaves. Free part of the bract a third of the
plexes forming the cone in 2 decussate pairs; upper size of the seed cone scale and not exceeding
fertile pair of scales spreading at maturity, with a long beyond the cone scale L. bidwillii
subapical bract tip; lower pair similar but smaller; col- 2b. Ultimate branchlets remain flattened, with
umella a small spike. Seeds 14 per cone, with 2 very facials less than half the size of laterals. Free
unequal wings. Seedlings with 2 cotyledons. part of the bract half the size of the seed cone
scale and exceeding beyond the cone scale
5 species. L. plumosa
3a. Leaves on ultimate branchlets of nearly equal
Distribution size. Lower pair of seed cone scales only slightly
smaller than upper pair L. chevalieri
New Zealand, New Caledonia. 3b. Leaves on ultimate branchlets very unequal in
size. Lower pair of seed cone scales much
Taxonomic notes smaller than upper pair 4
4a. Apex of facial leaves not reaching the next facial
Phylogenetic relationships among genera in leaf above, obtuse. Bract of lower, smaller seed
Cupressaceae based on DNA sequence data have cone scales twice as long as these
only more recently been analysed. Based on rbcL L. austrocaledonica
4b. Apex of facial leaves reaching the next facial keeled, with entire margins, bearing 4 small abaxial
leaf above, apiculate. Bract of lower, smaller yellow pollen sacs. Seed cones terminal on flattened
seed cone scales as long as these L. yateensis branchlets, developing within one growing season
to become thin woody, 1012 mm long. Bract-scale
complexes 68 35 mm, slightly rugose, spreading
Libocedrus austrocaledonica Brongn. & Gris, free parts of the bracts 57 1 mm on laterals and
Bull. Soc. Bot. France 18: 140. 1871. Stegocedrus 810 1.5 mm on facials. Columella a small spike
austrocaledonica (Brongn. & Gris) Doweld, Novosti ca. 1 mm long. Seeds 12, ovoid-oblong, slightly flat-
Sist. Vyssh. Rast. 33: 42. 2001. Type: New Caledonia: tened, with an acute to bifid apex, 56 2.5 mm, light
Grande Terre, Province Sud, Mont Humboldt, B. brown, with 2 opposite, thin membranous wings, the
522 Balansa 2503 (holotype P). smaller a narrow strip less than 1 mm wide, the larger
oval-oblong, 68 2.53 mm, yellowish brown.
Etymology
Distribution
The species epithet refers to its (predominant)
occurrence in the southern part of New Caledonia. New Caledonia: Province Sud, Province Nord (Mt.
Paoua).
Vernacular names TDWG codes: 60 NWC
No common names have been recorded for this New Caledonia: Province Sud (Mt. Humboldt, Mt.
species. Kouakou), Province Nord (Poindimi).
TDWG codes: 60 NWC
Description
Ecology
(Spreading) shrubs or small trees 25 m tall, mul-
tistemmed or sometimes monopodial, diam. ca. On slopes near the summits of some of the highest
10 cm. Bark rough and scaly, brown, exfoliating in peaks in usually steep terrain covered by a 25 m
thin irregular strips or plates. Branches numerous, tall maquis vegetation, in the contact zone between
ascending, forming a dense, bushy, often rounded schists and serpentines. The altitudinal range is from
crown. Foliage more or less frondose, forming dense 650 m to 1620 m a.s.l. These mountain summits
sprays, ultimate branchlets alternate to subopposite, receive high levels of precipitation.
Conservation to acute, appressed, partly covered at base by larger
25(6) 1.52 mm, divergent, bilaterally flattened,
This rare species is known from only three very slightly curved laterals with entire margins and
small populations on isolated mountain summits, free apices, leaves on older trees smaller and nearly
where growth is slow and regeneration poor. There is monomorphic; amphistomatic, stomata on facials
risk of wildfires even though two of the three popu- near base, on laterals in a small groove on upper-
lations are within floristic reserves. There is a likeli- side, in a broad, conspicuous band of irregularly but
hood of mining in the unprotected area. densely arranged stomata on underside; upperside
IUCN: CR [B1ab(iii)] green, underside with whitish green stomatal band.
Pollen cones terminal, solitary, subglobose to ovoid,
Uses 35 mm; microsporophylls decussate, 812, peltate, 525
with entire margins, bearing 4 abaxial yellow pollen
No uses are recorded of this species; it is grown sacs. Seed cones terminal on branchlets with weakly
in only a few botanic gardens, as young plants in dimorphic leaves of nearly equal size, developing
research greenhouses. within one growing season to become thin woody,
1218 mm long. Bract-scale complexes 1015 mm
long, with upper pair slightly recurved in distal half
Libocedrus plumosa (D. Don) Sarg., Silva N. Amer. and with an obtuse to truncate apex, subtended by
10: 134. 1896. Dacrydium plumosum D. Don, in the smaller (59 mm) acute pair, each with long
Lambert, Descr. Pinus, ed. 3, App.: . 1832. Type: excerted, reflexed and upcurved bract tips and finely
New Zealand: locality unknown, A. Cunningham rugose abaxial surface. Columella present in mature
330 (neotype K). Fig. 165 cones, a small conical spike 23 mm long. Seeds 24,
ovoid, flattened, with an acute apex, 35 mm long,
Etymology brown, with a whitish hilum and 2 opposite, thin
membranous wings; smaller wing a narrow strip less
The species epithet (Latin plumosus) means feath- than 1 mm wide; larger wing irregularly oval-oblong,
ery and refers to the appearance of the foliage. 68 34.5 mm, yellowish brown.
New Zealand cedar; kawaka (Maori) New Zealand: North Island, in South Island
restricted to the Tasman District.
Description TDWG codes: 51 NZN NZS
Manoao is the Maori name for at least one other and young plants distinct from adult leaves, acicu-
podocarp (Halocarpus kirkii) as well as for this lar, 610(12) mm long, bifacially flattened with a
one, between which the Maori presumably saw no midrib on both sides, straight or curved, stomata
distinction. in two distinct bands mainly on adaxial side; apex
acute. Branchlets with juvenile leaves may re-appear
Description among those with adult leaves. Intermediate phase
528 leaves often on sucker shoots, 45 mm long, dis-
See the species description. tichous, bilaterally flattened. Adult leaves spirally
arranged, imbricate and appressed, rhomboid in
Distribution appearance, 11.5 1 mm, keeled abaxially; apex
obtuse. Leaves amphistomatic, stomata conspicu-
As for the species. ous, scattered. Pollen cones terminal, sessile, 46
mm long, 22.5 mm wide; microsporophylls (6)8
12, rhombic to triangular, with minutely denticulate
Manoao colensoi (Hook.) Molloy, New Zealand upper margins and with two basal red pollen sacs.
J. Bot. 33: 196. 1995. Dacrydium colensoi Hook., Seed cones terminal, solitary or sometimes in pairs
Icon. Pl., n.s., 2: t. 548. 1843; Lagarostrobos colensoi on distally curved short branchlets, 34 mm long,
(Hook.) Quinn, Austral. J. Bot. 30 (3): 317. 1982. consisting of 25(6) fertile bracts. Seeds 13(5) per
Type: New Zealand: North Island, [High hills near cone, crowded, morphologically and topographi-
the eastern coast...], W. Colenso 27 (lectotype BM). cally erect, ca. 34 23 mm, rounded in cross-
Fig. 166 section, notched ar apex, dark purple to black, basal
half or more of seed enclosed in a swollen, fleshy,
Etymology yellowish green epimatium.
Distribution Conservation
Greek meta- = changed; i.e. a changed or trans- nearly so, mostly distichous, 515 cm long, 0.71 mm
formed Sequoia, denoting its close relationship and thick including decurrent leaf bases, glabrous, ter-
similarity. minating in small seasonal buds, deciduous. Leaves
opposite on foliage branchlets, the free part proxi-
Description mally constricted and twisted, merging with alter-
nating left and right trending decurrent bases,
530 See the species description. lamina linear, curved or nearly straight, spreading
at nearly right angles to branchlet, 815 mm (30
Distribution mm in young trees) 1.52.5 mm, generally longest
in middle section of branchlet, with parallel entire
As for the species. margins and a midrib, shortly tapering to obtuse or
mucronate apex. Leaves hypostomatic, stomata in
two broad bands of 48 lines divided by a narrow
Metasequoia glyptostroboides Hu & W. C. Cheng, raised midrib, leaf colour light green or sometimes
Bull. Fan Mem. Inst. Biol., ser. 2, 1 (2): 154. 1948. slightly glaucous green. Pollen cones numerous in
Type: China: Hubei, W Hubei, Modaoxi, [E spike-like shoot systems, opposite, ovoid, 56(10)
Szechuan, Wanhsien, Mo-tao-hsi], C. T. Hwa 2 2.54 mm when full grown; microsporophylls
(lectotype NF). Fig. 167, 168 1520, proximally helically arranged, but irregularly
attached in mature cones, peltate, with 3 abaxial pol-
Metasequoia glyptostroboides Hu & W. C. Cheng var. len sacs near lower margin. Seed cones terminal on
caespitosa Y. H. Long & Y. Wu, Bull. Bot. Res. North- 25 cm long, sparsely (scale-)leaved shoots, solitary,
East. Forest. Inst. 4 (1): 149. 1984. subglobose, barrel-shaped or fusiform when still
tightly closed, with nearly concentric grooves, glau-
Etymology cous green, maturing in one year to 1525(30)
1023 mm, parting the scales in the grooves, turning
The species epithet refers to its likeness with Glyp- dull brown or grey-brown. Bract-scale complexes
tostrobus. 1624(28), decussate, more or less peltate, dilated
into a transversely elliptic or broadly triangular disk
Vernacular names with a transverse indentation, 58 mm long, 815
mm wide, 24 mm thick, distally rugose, proximally
Dawn Redwood; shui shan (Chinese) moderately lignified, striate, yellowish to dark brown
at base. Seeds numerous, more or less inverted by
Description displacement, 46 45 mm including wings,
irregular, compressed, emarginate at apex, with two
Trees to 50 m tall (but most trees now under 35 m), broad, membranous yellowish or light brown wings
deciduous, monoecious; trunk monopodial, large encircling the seed.
trees often buttressed, d.b.h. to 2.2 m. Bark becom-
ing fissured, exposing purplish brown inner bark, Distribution
exfoliating in long, thin plates or strips. Branches
long, spreading to ascending, or foliage subpendu- Central China: Chongqing (Shizhu), Hubei
lous, forming a pyramidal crown in young trees, (Lichuan), Hunan (Longshan).
becoming broader and more open, with a rounded TDWG codes: 36 CHC-CQ CHC-HU CHC-SC
top, in older trees. Foliage branchlets opposite or CHS-HN
Ecology bottoms before farmers moved into the area only a
few centuries ago (Chii & Cooper, 1950). The sev-
The ecology of Metasequoia in undisturbed valley eral subpopulations are all reduced to a few to sev-
forests can only be reconstructed from palynologi- eral hundred mature trees, sometimes even a single
cal data and from clues obtained from field observa- tree, with little chance of natural generation due to
tions in the lower sections of some little side valleys changes in land use by a growing agrarian popula-
of the Shuishaba Valley, where relatively undis- tion. Secondary woodland in some narrow side val-
turbed stands occur on probably suboptimal sites. leys, where Metasequoia may occur along permanent
Remaining large trees in the valley proper are sur- streams, are under pressure of grazing and cutting
rounded by cultivated fields (mainly rice) and both of firewood. The mature, large trees have all been
native and introduced tree species, constituting in its declared protected but habitat protection is overall 531
most natural state a secondary vegetation. Though inadequate (Bartholomew et al., 1983; Fu & Jin, 1992;
some of the trees of Metasequoia have also been Wang & Guo, 2009), which means that the survival
planted, it is likely that the largest and oldest trees of this very interesting species in its natural habitat
in the valley are survivors. It is a riparian species is not guaranteed.
that occupies a habitat similar to that of Taxodium IUCN: EN [B1ab (iii, v)]
distichum; the remnant old trees may be the vestiges
of an extensive flood plain forest that existed before Uses
this valley was transformed to agriculture only a few
centuries ago. Away from the valley floor the trees In the past, trees of this species must have been used
are restricted to the moist bottoms of ravines and in for construction timber locally. Evidence of branch
contact with seepage water. The valley forest would cutting for firewood can be seen in the photograph
not have been pure Metasequoia, but mixed with of one of the earliest discovered trees, a picture that
angiosperms, among which were very likely species has been widely published. Its use is now prohibited
of Acer, Castanea, Populus, and Quercus, as well as in the Metasequoia area but the species has been
Liquidambar acalycina, Nyssa chinensis, Pterocarya widely planted, as an amenity or forest tree in China
hupehensis and other trees tolerant of periodic flood- and as an ornamental in many other countries with
ing. The soil is clay and sand derived from sand- temperate climates. This conifer is one of the most
stone, with slightly acid to neutral pH and a strongly remarkable success stories of introduced trees to
fluctuating but not deep water table. The climate is date; since its first introduction to the USA and
characterized by hot summers and cold winters. Europe in 1948 it has spread to almost every coun-
try with a temperate climate. Several cultivars have
Conservation been named, especially in the Netherlands. The phe-
notypic variation observed in planted trees obtained
This conservation flagship conifer species occurs in from early seed introductions may indicate genetic
a limited region in the border area of two Chinese diversity, but trees with somewhat stunted and con-
provinces and Chongqing Municipality (formerly E torted lower trunks may also be due to suboptimal
Sichuan province) in what is now intensively culti- growing (climatic?) conditions. It would seem that
vated and utilised countryside. It appears that at least warm and humid summer conditions are conducive
the older trees are relicts of a riparian Metasequoia to rapid growth, producing straight, erect trunks
forest, which may have stretched unbroken in valley and long branches.
Microbiota Kom., Bot. Mater. Gerb. Glavn. Bot. Sada RSFSR 4: 180. 1923. Type:
Microbiota decussata Kom. (Cupressaceae).
Greek: mikros- = small; kachrys = catkin, cone. minutely denticulate; dorsal side of leaves keeled
towards an obtuse, incurved apex. Stomata restricted
Description to adaxial side of leaves, subapical, hidden from
view. Pollen cones terminal, more or less recurved,
See the species description. 2.54 mm long, ovoid; microsporophylls helically
arranged, strongly incurved, enclosing two basal
534 Distribution pollen sacs containing bisaccate or trisaccate pollen.
Seed cones usually on different branching systems
As for the species. from pollen cones, terminal, maturing to 68 mm
long, ovoid, comprising of an axis with ca. 20 heli-
cally arranged, rounded bracts which become fleshy,
Microcachrys tetragona (Hook.) Hook. f., London slightly swollen and red at maturity. Ovules 1 per fer-
J. Bot. 4: 149. 1845. Athrotaxis tetragona Hook., Icon. tile scale, inverted; seeds broadly ovoid, 1.5 1.2 mm,
Pl., n.s., 2: t. 560. 1843, [Arthrotaxis]; Dacrydium slightly flattened when ripe, at base partly covered
tetragonum (Hook.) Parl., in Candolle, Prodr. 16 (2): by the epimatium.
496. 1868. Type: Australia: Tasmania, [V. D. L.], R.
C. Gunn [367] (lectotype K). Fig. 170 Distribution
Etymology Ecology
The species epithet describes the four-angled Microcachrys tetragona occurs in Tasmanian alpine
branchlets due to decussate phyllotaxis of the scale dwarf scrubland on exposed rock outcrops, usually
leaves. above 1000 m altitude. This prostrate little shrub
spreads over rocks and through low mossy or her-
Vernacular names baceous vegetation in wet areas with long, whip-
like running shoots capable of rooting and ramify-
Creeping pine, Strawberry pine ing towards open spaces, often over bare rock. It
is, despite being so low to the ground, wind polli-
Description nated as are all conifers; its tiny mulberry-like red
cones are succulent and undoubtedly eaten by ani-
Evergreen, monoecious or (temporarily) dioecious, mals (birds?), but these vectors of its seed dispersal
prostrate shrubs 1030 cm tall. Leading stems to ca. are not known. This dwarf conifer is often growing
1 m long, branching frequently. Foliage branches near other, taller conifers like Athrotaxis cupres-
numerous, spreading or ascending; whip shoots soides, Diselma archeri, Pherosphaera hookeriana,
elongated, creeping out over rocks; (pen)ultimate Podocarpus lawrencei, and various angiosperms,
branchlets with scale leaves quadrangular in cross- e.g. Nothofagus gunnii, Epacris serpyllifolia, Orites
section. Leaves on whip shoots mostly decussate, spp., Richea spp., Astelia alpina, Leptospermum
decurrent, lanceolate, to ca. 5 mm long, the distal ruprestre, etc.
part slightly spreading, with minutely denticulate
margins tapering to an acute apex. Leaves on lateral Conservation
branchlets strictly decussate, imbricate, appressed,
short triangular in appearance, 11.5 1 mm; margins IUCN: LC
Uses At Kew it was grown successfully in the Temperate
House and in the Himalayan House, but it has long
This extremely dwarfish conifer is extremely rare in since disappeared. In view of its natural habitat and
cultivation. Despite having been introduced to the climate, this species should be an interesting plant
Royal Botanic Gardens, Kew in 1862 by W. Archer, it for the rock garden in mild winter regions where
is still only known from a few botanical collections. frost is uncommon.
535
Nageia Gaertn., Fruct. Sem. Pl. 1: 191. 1788. Type: Nageia nagi (Thunb.) Kuntze
[Myrica nagi Thunb.] (Podocarpaceae).
Description Distribution
This species was named after the 19th century bota- Ecology
nist James Motley.
This species grows as tall as N. wallichiana in low-
Vernacular names land tropical rainforest from ca. 15 m to ca. 500 m
a.s.l. Nageia motleyi occurs usually on dry soil but
podo kebal musang (Peninsular Malaysia); kayu has also been found in peat swamps (ramin-peat
bawa, setebal (Sumatera); medang buloh (Sarawak). swamp) in Sarawak. It is a rare constituent of low-
land bindang-dipterocarp forest in Sarawak, but
Description is more often found in less tall evergreen forests on
podzolic sands, where it is scattered among numer-
Trees to 50 m tall; trunk to 1 m d.b.h., forming a ous angiosperm tree species and occasional conifers.
straight, clear bole. Bark hard and scaly, peeling in It apparently regenerates after disturbance and is
thin flakes, brown or reddish brown; inner bark also found in secondary forest.
fibrous, reddish. Crown with spreading branches,
becoming rounded. Foliage branchlets opposite (but Conservation
often one of the pair not developed) spreading rig-
idly, terete but ultimately more or less quadrangular Logging of Nageia spp. in lowland forests can involve
and often twisted, with grooves, glabrous; termi- this species or the much more widely distributed
nal buds with lanceolate scales extending to a fine species N. wallichiana; the ranges and ecology of
point. Leaves opposite or subopposite (towards end both species partly overlap in Peninsular Malaysia,
of branchlets often alternate), decussate or nearly in Sumatera and Borneo. Identification of logged trees
one plane on shaded branches; petiole twisted ca. to species is impossible if the foliage is no longer
90 at base, 23 mm long; leaf blade somewhat vari- available. It is likely that logging has affected this
able in size and shape, 25(7.5) 1.52.5 cm (larg- species more seriously than was previously realized.
erst on young plants and shaded leaves), lanceolate, IUCN: VU (A2c)
elliptic to obovate with obtuse, acute or acuminate
apex, coriaceous, with distinct parallel veins, dark Uses
green above, mid green below. Stomata on both
surfaces, in numerous, continuous or intermittent Nageia motleyi is a valuable timber tree, especially
lines, conspicuous on both sides. Pollen cones axil- where it grows into tall, straight trees with a long,
lary, solitary, sessile, 1.52 cm long, with a diam. of clear bole. It is traded as podocarp wood. Long tim-
56 mm; microsporophylls imbricate; apex acumi- ber is sawn into planks for construction (mainly
house building); other uses of the wood are plywood, ing rounded, or bushy. Foliage branchlets opposite
veneer, interior finishing, and furniture making. As (but often one of the pair not developed) spread-
it occurs within the general distribution of N. walli- ing rigidly, more or less tetragonal, often twisted,
chiana and develops into a similar tree, it is difficult with grooves in the edges, glabrous; terminal buds
to assess whether uses apply to the less frequently with narrowly lanceolate scales extending to a fine
encountered species or to the widespread species. point. Leaves opposite or subopposite (towards end
Nageia motleyi is probably not grown in cultivation. of branchlets often alternate), decussate or nearly in
one plane on shaded branches; petiole twisted ca.
Nageia nagi (Thunb.) Kuntze, Revis. Gen. Pl. 2: 798. 90 at base, 38(15) mm long; leaf blade variable in
1891. Myrica nagi Thunb., in Murray, Linn. Syst. size and shape, 29 0.73 cm (largerst on young
540 Veg., ed. 14: 884. Mai-Jun 1784; Decussocarpus nagi plants and shaded leaves), from narrowly elliptic
(Thunb.) de Laub., J. Arnold Arbor. 50: 357. 1969. to ovate-lanceolate or obovate with acute to obtuse
Type not designated. Fig. 172 or nearly truncate apex, coriaceous, with indistinct
parallel veins, dark green above, pale green below.
Podocarpus formosensis Dummer, Gard. Chron., Stomata only on lower surface, in numerous, inter-
ser. 3, 52: 295. 1912; Decussocarpus nagi (Thunb.) de mittent lines, usually conspicuous but sometimes
Laub. var. formosensis (Dummer) Silba, Phytologia poorly visible. Pollen cones axillary, in clusters of
58: 366. 1985; Nageia formosensis (Dummer) C. N. up to 10 or sometimes solitary, (nearly) sessile, vari-
Page, Notes Roy. Bot. Gard. Edinburgh 45: 382. able in length from 0.52 cm, becoming cylindrical
1989; Nageia nagi (Thunb.) Kuntze var. formosensis at full length, with a diam. of 23 mm; microspo-
(Dummer) Silba, Phytologia 68: 38. 1990. rophylls imbricate, apiculate, with two pollen sacs.
Podocarpus nankoensis Hayata, [Icon. Pl. Formos. 6, Seed cones axillary, solitary and long pedunculate,
Suppl.: 74. 1917, nomen] Icon. Pl. Formos. 7: 39. 1918; or rarely in pairs, with a few deciduous bracts near
Nageia nankoensis (Hayata) R. R. Mill, Novon 9 (1): base of fertile branchlet and some remaining dried
77. 1999. bracts subtending a single seed enclosed in a green,
Podocarpus nagi (Thunb.) Pilg. var. koshunensis when ripe dark purple epimatium, usually bloomed
Kaneh., Trans. Nat. Hist. Soc. Formosa 21: 145. 1931; white, the whole (sub)globose and 1015 mm diam.
Podocarpus formosensis Dummer var. koshuensis when succulent, wrinkled and dark brown when dry
(Kaneh.) Merr. & Yamam., J. Soc. Trop. Agric. 7: with a loose seed inside.
142. 1935; Podocarpus koshunensis (Kaneh.) Kaneh.,
Formosan Trees, ed. 2: 36. 1936; Nageia nagi (Thunb.) Taxonomic notes
Kuntze var. koshuensis (Kaneh.) D. Z. Fu, Acta
Phytotax. Sin. 30 (6): 524. 1992. Nageia formosensis (Dummer) C. N. Page was listed
in A World Checklist and Bibliography of Conifers
Etymology (Farjon, 1998, [2001]) as an accepted species. It was
by some considered a mere variety of N. nagi, while
The species epithet refers to the Japanese vernacular other sources suggest that perhaps there are no
name as recorded by Carl Peter Thunberg. fewer than three distinct species in Taiwan: Nageia
formosensis, N. nankoensis (Hayata) R. R. Mill and
Vernacular names N. fleuryi (Hickel) de Laub. Given the variability I
have observed in specimens considered to belong
naki (Japanese); zhu bai (Chinese) to N. nagi from mainland China, Taiwan and Japan
(some of which came from planted trees), a broader
Description concept of N. nagi is here adopted (as in Flora of
China 4, 1999), until it can be clearly demonstrated
Trees or shrubs to 20 m tall; trunk to 50 cm d.b.h. that there are distinct and sufficient character state
Bark scaly, peeling in small, thin flakes, red- differences in the Taiwanese trees that merit the rec-
dish or purplish brown, weathering grey. Crown ognition of one or more separate species, or even
with ascending or spreading branches, becom- varieties of N. nagi.
Distribution based on intimate knowledge of the forest flora are
lacking for most of mainland China.
S China: Anhui, Fujian, Guangdong, Guangxi, IUCN: NT
Hainan, Hunan, Jiangxi Zhejiang; Japan: Kyushu,
Nansei-Shoto [Ryukyu Is.], S Honshu, Shikoku; Uses
Taiwan.
TDWG codes: 36 CHC-HU CHH CHS 38 JAP-HN Nageia nagi is a valuable timber tree, but its most
JAP-KY JAP-SH NNS TAI common use is as an amenity tree in China and
Japan, where it is found in many of the climatically
Ecology milder parts of these countries planted in gardens,
parks, sanctuaries, and even as street trees. It is also 541
Nageia nagi occurs in mixed mesophytic evergreen popular as a tree for bonsai cultivation. It is much
forest and mixed mesophytic deciduous forest less commomly planted in Europe, the USA and
(Wang, 1961). It occurs in hills and low mountains New Zealand, where it is almost restricted to botani-
from about 200 m to 1200 m a.s.l. In evergreen oak cal collections.
forest it is one of several shade tolerant conifers that
may occur under canopy or take advantage of small
gaps to break through: Taxus chinensis, Cephalotaxus Nageia wallichiana (Presl) Kuntze, Revis. Gen.
fortunei, Keteleeria fortunei, and Fokienia hodginsii Pl. 2: 800. 1891. Podocarpus wallichianus C. Presl,
are the most common of these. Besides Castanopsis Abh. Knigl. Bhm. Ges. Wiss., ser. 5, 3: 540. 1846;
spp. and Quercus spp. (the oaks), numerous angio- Decussocarpus wallichianus (C. Presl) de Laub., J.
sperm trees occur in these forests or forest rem- Arnold Arbor. 50: 349. 1969; Podocarpus latifolius
nants. In Taiwan and southern Japan the coniferous Wall., Pl. Asiat. Rar. 1 (2): 26, t. 30. 1830, non Mirb.
element of this vegetation is more dominant, with (1825); Nageia latifolia (Wall.) Gordon, Pinetum:
Pseudotsuga sinensis or P. japonica and Tsuga siebol- 138. 1858. Type: India: Pundu Mts., Mt. Sillet, [in
dii often added to the mixture. In forest or woodland montibus Punduae], N. Wallich 6050 (holotype
on drier mountain slopes N. nagi tends to follow K-W). Fig. 173, 174
streams, but it is known to regenerate in more open
thickets after forest disturbance. Etymology
Ecology
Callitropsis R. H. Compton, J. Linn. Soc., Bot. 45: strips, grey. Branches spreading wide or ascending,
432. 1922, non Oerst. (1864). Type: Callitropsis arau- higher order branches assurgent or erect, forming
carioides R. H. Compton [Neocallitropsis pancheri a conical or more often a rounded or flat-topped
(Carrire) de Laub.] open candelabra crown. Foliage branchlets in
dense tufts of 2030, crowded at the terminal 1015
Latin: neo- = new; Callitropsis (genus name) means cm of main branches, assurgent or erect, sparsely
similar to Callitris; the prefix is here used to avoid a branched or unbranched, to 20 cm long, 510(12) 543
later homonym. mm wide, densely covered with imbricate leaves,
mostly determinate and deciduous, some continu-
Description ous. Leaves in alternate whorls of 4, whorls turned
45 degrees around shoot axis relative to the previ-
See the species description. ous whorl, seemingly in 8 rows, short decurrent or
nearly sessile with broad base, lanceolate, incurved,
Distribution thick, coriaceous, rigid, keeled, 615 1.82.5 mm;
margins minutely serrate; apex acute-pungent;
As for the species. stomata in two bands abaxially, in more scattered
rows adaxially; leaf colour yellowish green or green.
Pollen cones terminal, 1012 67 mm, subglobose
Neocallitropsis pancheri (Carrire) de Laub., Fl. to ovoid; microsporophylls in 34 alternating whorls
Nouv. Caldonie Dpend. 4: 161. 1972. Eutacta of 4, 36 2 mm, peltate-rhombic with a long, acute
pancheri Carrire, Trait Gn. Conif., ed. 2, 2: 864 or acuminate apex, bearing (2)614 abaxial, small
[615]. 1867. Type: New Caledonia: Grande Terre, pollen sacs. Seed cones terminal on short branch-
Province Sud, Montagnes de Yat, E. Vieillard 1274 lets, maturing within a year to an opened cone up to
(holotype P). Fig. 175, 176 15 mm long with spreading bract-scale complexes.
Mature bract-scales in two alternating whorls of 4,
Callitropsis araucarioides R. H. Compton, J. Linn. with lower scales slightly larger than upper ones,
Soc., Bot. 45: 432. 1922; Neocallitropsis araucarioides 1012 3 mm and 810 2 mm, subulate to linear,
(R. H. Compton) Florin, Palaeontographica B 85 similar to mature foliage leaves but with adaxial
(18): 590. 1944. thickening, leaving margins of outer bracts and api-
ces unaltered but reflexed, rostrate; margins of thick-
Etymology ened tissue papillose. Columella short pyramidal,
1.5 mm tall. Seeds 12(4) per cone, angular-ovoid,
The species epithet commemorates the French bota- acutish, 67 2.53.5 mm, light brown, with 2(3)
nist Jean A. I. Pancher (18141877). marginal 0.50.7 mm wide wings.
No common names have been recorded for this New Caledonia: Province Sud (SE part), one loca-
species. tion in Province Nord (Mt. Paoua).
TDWG codes: 60 NWC
Description
Ecology
Shrubs or small evergreen trees to 46(10) m tall,
multistemmed or monopodial, stem to 3050 cm In scrubland (maquis minier) on ultramafic rock
diam. Bark on large stems exfoliating in narrow (serpentine, cuirasse maquis), following stream
courses and along lower mountain ridges up to about unsustainable exploitation of the wood of this slow
950 m a.s.l., rarely higher. This species is often asso- growing conifer for oil extraction, but this use has
ciated along streams with the podocarps Dacrydium now virtually ceased. No assessment of any decline
araucarioides, D. guillauminii, and Dacrycarpus due to this exploitation has come to the attention of
vieillardii, elsewhere with Agathis ovata, Callitris the Conifer Specialist Group of IUCN-SSC, yet the
neocaledonica, Podocarpus novae-caledoniae, and slow growth of this species makes it likely that this
angiosperms, including many shrubs as well as has occurred. A large population covering ca. 122 ha
Cyperaceae. Several of the trees and shrubs also is protected in the Montagne des Sources (Parq de
develop the candelabra-like crowns typical for low la Rivire Bleue) and smaller populations are under
trees and shrubs in this environment, but grow more protection at the Chte de la Madeleine (17.5 ha) and
544 solitary than Neocallitropsis, which is strongly gre- in the Plaine des Lacs (3.5 ha). Despite this protec-
garious. The climate is tropical with abundant rain- tion, continued decline and fragmentation of popu-
fall through most of the year. lations are projected.
IUCN: EN [A2c, B1ab(iii)+2ab(iii)]
Conservation
Uses
The habitat of this species is extremely susceptible
to fire and wildfires are a widespread hazard in New The resin in the wood of this species is used to make
Caledonia. With less than 10 populations known in an oil extract, which is sold as Araucaria oil; this is
the southern mountains and one, recently discov- now only locally traded. It is also locally used as an
ered, in the northern part of Grande Terre, con- ornamental shrub, but does not appear to have been
servation issues are apparent. In the past, there was taken into cultivation outside New Caledonia.
A HA N DB O OK OF T H E WOR L D S C ON I F E R S
Giant Sequoia (Sequoiadendron giganteum) Drawing by Aljos Farjon
A HA N DB O OK OF T H E WOR L D S C ON I F E R S
by
AL JO S FA R JON
Volum e II
LEIDEN-BOSTON
2017
This book is printed on acid-free paper.
Library of Congress Cataloging-in-Publication Data
The Library of Congress Cataloging-in-Publication Data is available from the Publisher.
front cover:
Cunninghamia konishii foliage
back cover:
top left: Pinus wallichiana seed cones
bottom right: Cephalotaxus haringtonii ripe seeds
Nothotsuga Hu ex C. N. Page, Notes Roy. Bot. Gard. Edinburgh 45: 390. 1989. Type:
Nothotsuga longibracteata (W. C. Cheng) Hu ex C. N. Page [Tsuga longibracteata
W. C. Cheng] (Pinaceae).
Greek: nothos = bastard, base-born; i.e. a (putative) ovoid, acutish at apex, 24mm long, not resinous,
hybrid between Tsuga and an unnamed genus. shining brown. Leaves more or less pectinate, on
small lateral shoots in false whorls on emergence,
Description otherwise remote, with oblique, twisted petiole,
1.12.4cm long, 12(2.5)mm wide, linear, abruptly
See the species description. tapering at both ends; apex acutish, faintly grooved 549
above, slightly flattened; margins entire; stomata
Distribution 510 lines along the median groove above, two
bands separated by a faint midrib below (abaxial
As for the species. side); leaf colour dark glossy green, stomatal bands
whitish green. Pollen cones subterminal on small
lateral shoots, clustered in umbels from a single
Nothotsuga longibracteata (W. C. Cheng) Hu ex bud, pedunculate, pendant, 510mm long, yellow-
C. N. Page, Notes Roy. Bot. Gard. Edinburgh 45: ish brown (in sicco). Seed cones lateral, or subtermi-
390. 1989. Tsuga longibracteata W. C. Cheng, Contr. nal on small shoots, more or less erect on 510mm
Biol. Lab. Sci. Soc. China, Sect. Bot. 7 (1): 1. 1932. long peduncles, ovoid oblong to cylindrical, with
Type: China: Guizhou, Yinjiang Tujiazu Miaozu obtuse truncate apex, (2)2.55(5.8)cm long, 1.5
Zizhix, Fanjing Shan, [Yinkiang], Y. Tsiang 7712 2.5(3)cm wide with opened scales, purplish or red
(holotype NAS). Pl. 22, Fig. 177, 178 when immature, ripening to dark brown; old cones
persisting several years, breaking off at peduncle,
Etymology or sometimes disintegrating; cone rachis blackish
brown. Seed scales 2030, suborbicular to broadly
The species epithet describes the relatively long peltate-auriculate, convex, opening slightly or very
bracts on the seed cone. wide (reflexed), 12.2 1.22.5cm at mid-cone;
abaxial surface sparsely pubescent when imma-
Vernacular names ture, soon glabrous, finely striated; upper margin
rounded, entire or erose; base short pedicellate.
Bristlecone Hemlock; changbao tieshan (Chinese) Bracts subspathulate, erose-denticulate at acute
apex, 718mm long, straight, not exserted beyond
Description margins of more distal seed scales. Seeds ovoid, 4
2.53mm, brown; seed wings ovate-oblong, 712
Trees to 30m tall, d.b.h. to 11.2m; trunk monopo- 56mm, reddish brown.
dial, often forked or multi stemmed; bark soon flak-
ing, rough and scaly, brownish grey. Branches long, Taxonomic notes
heavy, curved, ascending or spreading; crown of
young trees conical, with a drooping leader, of old This taxon was first described as a species of Tsuga
trees open and irregular or dense and flat topped. by the well-known Chinese botanist W. C. Cheng in
Branchlets slender, firm, not drooping, brown- 1932. Hu (1951) proposed a separate genus Nothotsuga
ish yellow to brown, in 2nd or 3rd year grey; mostly for this species, but failed to give a Latin descrip-
glabrous, rarely minutely pubescent, with weakly tion; the genus name was then validated by Page
developed pulvini; on 23 year old twigs develop (1989). French botanists in the School of Gaussen at
numerous small, lateral, leaved shoots (515mm), Toulouse proposed a generic hybrid origin between
which do not extend much further. Vegetative buds Keteleeria and Tsuga, but gave no evidence for this
550
Plate 22. Nothotsuga longibracteata. 1. Habit of trees. 2. Branch with foliage. 3, 4. Seed cones. 5. Seed cone
scales. 6. Bract. 7, 8. Leaves. 9. Seeds. 10. Short shoot with leaves. 11. Pollen cones.
and applied an illegitimate name. Chinese botanists sinensis, Nyssa sinensis, Acer angustilobium, Davidia
(e.g. Flora of China 4: 3940, 1999) do not recognize involucrata, Sorbus spp., etc. In the evergreen broad-
its status as a distinct genus, but there are several leaved forest formation there are stands of pure
distinctive characters in both male and female cones Nothotsuga longibracteata and Tsuga chinensis. Pinus
not shared by other species of Tsuga in Asia or North massoniana or P. fenzeliana (syn. P. kwangtungensis)
America that appear to justify generic recognition. locally dominate the general canopy of broad-leaved
Its phylogenetic position based on both morpholog- trees on poorer sites, where N. longibracteata is also
ical and DNA data confirms this taxonomy. Despite concentrated.
its name, there is no evidence that this taxon is of
hybrid origin. Conservation
551
Distribution This species has been considered to be Endangered,
because it is very rare despite its relatively wide
China: Fujian, N Guangdong, NE Guangxi, NE distribution. Large scale logging has depleted the
Guizhou, SW Hunan, SE Jiangxi. number of trees to an unquantified extent (Fu & Jin,
TDWG codes: 36 CHC-GZ CHS-FJ CHS-GD CHS-GX 1992) and substantial parts of forest with this species
CHS-HN CHS-JX must have gone, especially at lower elevations. In
spite of the rarity of the species throughout its range,
Ecology it is not felt that the population reduction could have
exceeded 70% or even 50% over the past three gen-
Nothotsuga longibracteata occurs on low to medium erations which would prevent a category of threat
high mountains, at elevations between 300 and from being applied.
2300m a.s.l. It grows on both red and yellow earth. IUCN: NT
The climate is humid and warm-temperate to wet
and cool, with annual precipitation between 1000 Uses
2000mm. The species occurs in two forest forma-
tions (Wang, 1961). In the evergreen broad-leaved In China this species is considered to be a desirable
forest formation mostly with sclerophyllous broad- forest tree suitable for afforestation. Its use as a tim-
leaved trees such as Castanopsis spp., Lithocarpus ber tree must be limited due to its rarity. It is not in
spp., Quercus spp., and with Fokienia hodginsii; in general cultivation outside China and rare in botani-
the deciduous mixed mesophytic forest at higher cal collections.
elevations with Fagus longipetiolata, Tetracentron
Papuacedrus H. L. Li, J. Arnold Arbor. 34: 25. 1953. Type: Papuacedrus papuana
(F. Muell.) H. L. Li [Libocedrus papuana F. Muell.] (Cupressaceae).
Papua = New Guinea; Cedrus is the classical name or decussate, strongly dimorphic, imbricate; facials
for (true) cedars. much smaller than laterals, 1mm on old, mature
foliage, up to 8mm long on whip shoots of young
Description plants, rhombic to lanceolate in appearance, cari-
nate, cuspidate; laterals bilaterally flattened, 23mm
See the species description. long on old, mature foliage, up to 20mm long on
552 whip shoots of young plants, lanceolate to oblong;
Distribution margins entire, with incurved, appressed or free,
obtuse or acute apex; few stomata on upperside and
As for the species. 2 broad primary stomatal bands on underside; gland
absent or inconspicuous at base of facials; leaf colour
lustrous olive green to dark green, with glaucous
Papuacedrus papuana (F. Muell.) H. L. Li, J. Arnold green stomatal bands. Pollen cones terminal, soli-
Arbor. 34: 25. 1953. tary, cylindrical, 625 23mm; microsporophylls
830, decussate or in whorls of 4, peltate, with acute
Etymology or rounded apex, bearing (2)4(6) abaxial pollen
sacs near lower margin. Seed cones (sub)terminal, on
The species epithet refers to Papua, a collective name 210mm long branchlets with rhombic, acute scale
for the island of New Guinea and the Australasian leaves, thin woody, 818 58mm when closed,
people who were its first settlers. changing from green to glaucous green to brown or
dark blackish brown. Bract-scale complexes consist-
Vernacular names ing of 2 decussate pairs; lower pair 47 25mm,
curved, widest at base; upper pair much larger, 717
De Laubenfels, in Flora Malesiana 1, 10 (3): 446 37mm, elliptic, widest in the middle section
(1988), cites numerous local names applied in New where the recurved, acute bract tip emerges, distal
Guinea to this species. part slightly reflexed, truncate or obtuse, rugose with
grooves radiating from bract tip abaxially, smooth or
Description striate adaxially, with whitish seed scars near base.
Seeds 24, angular-ovoid or oblique, 25 13mm,
Trees to 20(50)m tall, evergreen, monoecious, red-brown with a whitish hilum; wings 2 on opposite
often appearing dioecious; trunk normally mono- sides, thin membranous; largest wing 47 25mm;
podial, slender, to 60cm d.b.h. Bark often spirally smallest wing often reduced to a strip 12mm wide,
twisted, exfoliating in shaggy scales or strips; outer translucent yellowish brown.
bark light brown, weathering grey. Branches short
and spreading in sheltered (younger) trees, long, Taxonomic notes
ascending to nearly erect in high montane forest,
forming conical to pyramidal crowns, or umbel- The morphological differences in the leaves between
late, flat-topped crowns at higher altitudes. Foliage Papuacedrus papuana var. papuana and P. papuana
branches mostly in flattened sprays, denser and var. arfakensis are minor and disappear with the
assurgent in older and exposed crowns, branching maturation of the foliage. As in many other conifers,
distichous, frondose, ultimate branchlets gradu- leaves in a juvenile stage differ markedly in shape and
ally shorter, entirely covered with flattened leaves. size from leaves in an adult stage and, as in this case,
Leaves on lateral branchlets scale-like, in whorls of 4 there are often transitional forms as well. The ranges
of both varieties overlap, but var. arfakensis only is generally high but seasonal, with up to 4000mm
occurs in the western parts of New Guinea and is per year on the highest slopes.
presumably the only variety in the Moluccas, where
the species remains undercollected (Johns, 1995). Uses
From herbarium specimens with adult-type foliage
and without pollen cones it becomes impossible to The timber of this species is widely used for con-
determine whether the sample represents either one struction (mainly building of houses in villages); in
of the varieties. Van Royen (1979), perhaps with this some areas the fibrous bark is used for roof cover-
difficulty in mind, recognized one species without ing and insulating house walls. It is rarely cultivated,
any infraspecific distinctions. although individual trees may be planted at sing-
sing or dance grounds of villages (Johns, 1995). In 553
Distribution horticulture it is only known from a few botanic
gardens.
Malesia: Maluku [Moluccas]; Papuasia: New Guinea.
TDWG codes: 42 MOL 43 NWG-IJ NWG-PN 2 varieties are recognized:
Ecology
Papuacedrus papuana (F. Muell.) H. L. Li var.
This species, as it occurs along a substantial altitu- papuana. Libocedrus papuana F. Muell., Trans.
dinal gradient, is present in different forest zones Roy. Soc. Victoria 1 (2): 32. 1889. Type: Papua
from montane tropical rainforest to subalpine New Guinea: Owen Stanley Range, Mt. Knutsford,
scrubland. The altitudinal range is (620)900 [probably Owalama Range], W. MacGregor 286
3600(3800)m a.s.l.; its greatest extent and abun- (holotype MEL). Fig. 179, 180
dance is reached in the mossy cloud forest zone
from ca. 1500m to the tree line. In the lower mon- Description
tane rainforest it is a scattered emergent associated
with angiosperms such as Casuarina, Castanopsis, Lateral pair of transitional leaves on young plants
Cinnamomum, Engelhardtia, Halfordia, Lithocarpus, spreading widely from base up to 6mm at the far-
Schizomeria, and Xanthostemon; in higher montane thest point below apex, falcately curved; apex turned
forest it becomes more prevalent with Nothofagus, upwards or slightly recurved; intermediate leaves
Cryptocarya, and Eugenia. It can also form coni- similar but less widely spreading. Pollen cones
fer dominated forest, occasionally with Araucaria 615mm long, with up to 16 decussate or whorled
cunninghamii var. papuana, but more often with microsporophylls.
Dacrydium spp., Phyllocladus hypophyllus, and
Podocarpus spp. These small conifer forests are often Distribution
surrounded by fire-induced grasslands (Imperata
cylindrica) with a mantle zone of ericaceous shrubs, New Guinea
or by tree fern grassland. Many of these conifers are TDWG codes: 43 NWG-IJ NWG-PN
often present in lower densities in the lower montane
rain forests as well. At lower altitudes Papuacedrus Conservation
grows mostly on basic soils. In mossy forest, along
high mountain streams and in subalpine scrubland Despite logging activities in large parts of its range
the trees are stunted and the soils are often acidic and that have been associated with human habitation for
water-logged; at around 3000m or higher swamps a very long time, this variety is still abundant and
are often surounded by Papuacedrus-Phyllocladus- not considered threatened with extinction.
Podocarpus woodland with tussocks of Gahnia IUCN: LC
dominant in the understorey (Johns, 1995). Rainfall
Distribution
Parasitaxus de Laub., Fl. Nouv. Caldonie Dpend. 4: 44. 1972. Type: Parasitaxus
usta (Vieill.) de Laub. [Dacrydium ustum Vieill.] (Podocarpaceae).
Latin: parasitus = a parasite; Taxus is the classical scaly with leaves, purple. Leaves scale-like, spirally
Latin name for yew. [While now understood as a arranged, imbricate and decurrent, 23mm long,
distinct family, podocarps were formerly classified 12mm wide (on older branches larger and being
with Taxaceae]. forced apart by the thickening branch), broadly lan-
ceolate to more or less triangular; apex appressed or
Description free, obtuse or acute. Stomata on both sides of leaves,
conspicuous and scattered on abaxial side. Pollen 555
See the species description. cones usually on the same branching systems as seed
cones, terminal, solitary, more or less oval, 33.5mm
Distribution long, 1.52mm wide; microsporophylls 813, imbri-
cate, broadly triangular with erose upper margin,
As for the species. terminating in an incurved apex, with two basal pol-
len sacs. Fertile ovulate branchlets numerous, erect.
Seed cones mostly terminal, some lateral, consisting
Parasitaxus usta (Vieill.) de Laub., Fl. Nouv. of 36 spreading narrow and apically incurved red
Caldonie Dpend. 4: 45. 1972, [ustus]. bracts of which the distal 12 subtend short pedun-
Dacrydium ustum Vieill., Ann. Sci. Nat. Bot., sr. 4, culate fertile scales bearing a single terminal and
16: 56. 1861; Podocarpus ustus (Vieill.) Brongn. & inverted ovule. Young seeds ovoid-oblong with an
Gris, Bull. Soc. Bot. France 13: 426. 1866; Nageia apical crest, when mature completely surrounded
usta (Vieill.) Kuntze, Revis. Gen. Pl. 2: 800. 1891. by a globose, hard, glaucous white epimatium, the
Type: New Caledonia: Grande Terre, Province Sud, whole 34mm diam.
Poila, (mountains around Poila), E. Vieillard 1267
(holotype P). Fig. 181, 182 Taxonomic notes
Conservation
Pherosphaera W. Archer, Hookers J. Bot. Kew Gard. Misc. 2: 52. 1850, p.p., quoad
descr. foem. Type: Pherosphaera hookeriana W. Archer (Podocarpaceae).
Microstrobos J. Garden & L. A. S. Johnson, Contr. Pherosphaera fitzgeraldii (F. Muell.) Hook. f.,
New South Wales Natl. Herb. 1 (6): 315. 1951. Type: Hookers Icon. Pl. 4: t. 1383. 1882. Dacrydium
Microstrobos fitzgeraldii (F. Muell.) J. Garden & fitzgeraldii F. Muell., Fragm. 11: 102. 1880;
L. A. S. Johnson Microstrobos fitzgeraldii (F. Muell.) J. Garden &
L. A. S. Johnson, Contr. New South Wales Natl.
Greek: phero = to bear; sphaira = ball, globe; refer- Herb. 1 (6): 316. 1951. Type: Australia, New South
ring to the shape of the seed cones. Wales, Blue Mountains, R. Fitzgerald s.n., 1880, 557
1881 (syntypes, ?MEL, n.v.).
Description
Etymology
Dioecious evergreen shrubs. Resin cavities in leaves.
Bark smooth or rough and scaly. Branches spreading This species was named by Ferdinand von Mueller
and rigid or pendulous and creeping. Adult and juve- after R. Fitzgerald, who presented him with the first
nile leaves similar, spirally arranged, 24(6)mm botanical specimens.
long and mostly scale-like. Stomata on adaxial side,
hidden from view. Pollen cones terminal, globose Vernacular names
to ovoid, 26mm long; microsporophylls (6)815,
spirally arranged, with two basal pollen sacs; pol- Dwarf mountain pine
len with 23 air sacs. Seed cones terminal on small
branchlets, more or less globular, 24mm long, Description
with (3)58 spreading, fertile scales, not fleshy at
maturity. Mature seeds usually 14 per cone, solitary Ascending, erect or drooping shrub to 1m tall and
and basal on adaxial side of a fertile scale (bract), 2m wide. Bark smooth, brown, weathering grey.
exposed, erect, ovoid with a slightly constricted and Branches slender, drooping to pendulous or strag-
truncated apex. gling over rocks; foliage branches very lax and slen-
der, irregularly branched two to four times. Leaves
2 species. monomorphic, spirally arranged, decurrent at base,
the free part spreading, 24mm long (varying in
Distribution length on a single branchlet), ca. 1mm wide (wider at
base, narrower at apex), subulate, apically incurved,
Australia: New South Wales, Tasmania. keeled on abaxial side, convex on adaxial side; apex
obtuse or apiculate; leaf colour dark olive green,
Key to the species of Pherosphaera whitish on adaxial side where stomata are situated.
Pollen cones terminal, solitary, globose maturing
Branches drooping to pendulous; foliage to ovoid, 46 3mm; microsporophylls 1015, spi-
branches very lax and slender; leaves 24 rally arranged, ovate-oblong with erose-denticulate
1mm P. fitzgeraldii margin and two basal pollen sacs. Seed cones ter-
Branches spreading, contorted; foliage branches minal and solitary on erect branchlets, more or less
assurgent or spreading; leaves 1 1mm globular, 24mm long, with 48 fertile, spreading,
P. hookeriana broadly lanceolate to ovate bracts. Mature seeds usu-
ally 13 per cone, solitary and basal on adaxial side
of a fertile scale (bract), exposed, erect, ovoid with
a slightly constricted and truncated apex, ca. 1
0.7mm, dull brown.
This species is extremely rare and has suffered a sub- Mount Mawson pine
stantial decline. Baker & Smith (1910) wrote: at the
base of most of the chief falls on the Blue Mountains Description
but it is now known to grow only on a few cliffs in
the Wentworth Falls and Katoomba Falls, an extent Erect, densely branched shrub to 2.5m tall. Bark
of occurrence ca. 9 km long. A survey in 1988 on larger stems rough, exfoliating with small scales,
reported 455 individual plants in 7 populations. Five brown weathering blackish grey. Branches spreading,
of these populations were on government land (Blue contorted; foliage branches assurgent or spreading,
Mountains National Park), two on private land. divided up to four times until very small; (pen)ulti-
Recruitment is very low or almost negligible and a mate branchlets 11.5mm thick, forming dense and
slow decline has been observed directly over several stiff tufts of foliage. Leaves spirally arranged, scale-
years. Threats are or have been urban development, like, in seedlings and young plants short lanceo-
water pollution, habitat degradation, and competi- late, 22.5 1mm, largely free but incurved, keeled,
tion from invasive species (Hedera helix, Rubus sp.), replaced higher up the plant by smaller leaves; in
as well as native shrubs and trees. mature plants all imbricate and closely appressed,
IUCN: CR [B1ab (iii)] broadly triangular, mostly ca. 1 1mm, on whip
shoots to 2.5 1.5mm, on older, thicker branchlets
Uses to 2 2mm, concave with a blunt keel; margins
minutely serrate; apex obtuse; leaf colour yellowish
Dwarf mountain pine is rare in cultivation, but green. Stomata on adaxial side, hidden from view.
makes an attractive rock-dwelling small shrub. It is Pollen cones numerous, terminal, globose, ca. 2mm
presently mostly confined to botanic gardens, but diam. colour red-brown at anthesis; microsporo-
phylls (6)812, spirally arranged, rounded with in winter. There is no extended period of snow cover
serrulate margins and two basal pollen sacs. Seed as the climate is extremely oceanic. Bedrocks are
cones terminal on distally down-curved branchlets, acidic granites, gabbro, and gneiss and the waters
more or less globular, 34mm long, with (3)58 have a low pH of 4.55 on average. This species
spreading, fertile, ovate and concave scales with an is often associated with Athrotaxis cupressoides,
acute apex, browning at maturity. Mature seeds usu- Microcachrys tetragona and, usually on somewhat
ally 14 per cone, solitary and basal on adaxial side drier sites, with Diselma archeri; frequent angio-
of a fertile scale (bract), exposed, erect, ovoid with sperms are Nothofagus gunneri, Richea pandanifolia,
a slightly constricted and truncated apex, ca. 1.3 R. scoparia, and Eucalyptus coccifera, while cushion
1mm, lustrous brown. forming peat mosses (Sphagnum) cover the ground
in many places. 559
Distribution
Conservation
Australia: Tasmania (central and western moun
-
tains). IUCN: NT
TDWG codes: 50 TAS
Uses
Ecology
This shrubby species is rare in cultivation, being
Pherosphaera hookeriana occurs in the subalpine grown in some specialist nurseries and often con-
regions of the Tasmanian highlands, usually above fused with Diselma archeri (Cupressaceae). It
1000m a.s.l. It is frequent in wet moors and often should be hardy in areas with mild and wet winters
fringes the lakes and tarns that are numerous in where frost is light. It will make a suitable shrub
these mountains. Precipitation is high and occurs for rock gardens or for pot-grown balcony or patio
year-round; temperatures are cool with sleet and evergreens.
snow falling in most months of the year, but mostly
Phyllocladus Rich. ex Mirb., Mm. Mus. Hist. Nat. 13: 48. 1825 (nom. cons.). Type:
Phyllocladus aspleniifolius (Labill.) Hook. f. [Phyllocladus billardieri Mirb. (nom.
illeg.) (Podocarpus aspleniifolius Labill.)] (Phyllocladaceae).
Brownetera Rich. ex Tratt., Ann. Mus. Natl. Hist. Nat. and subtended by a filmy white aril leaving the apical
(Paris) 16: 299. 1810. Type: Brownetera aspleniifo- part free. Seedlings with 2 cotyledons.
lia (Labill.) Tratt. [Podocarpus aspleniifolius Labill.]
Thalamia Spreng., Anleit. Kennt. Gewchse, ed. 4 species.
2, 2: 218. 1817. Type: Thalamia aspleniifolia (Labill.)
560 Spreng. [Podocarpus aspleniifolius Labill.] Distribution
Greek: phyllos = leaf; klados = branch, shoot; in Malesia: Borneo, Sulawesi, Philippines, Maluku
botanical Latin: phyllocladium = a leaf-like branch [Moluccas], New Guinea; Australia: Tasmania; SW
or shoot. Pacific: New Zealand.
Shrubs or trees, usually dioecious, occasionally mon- Reproductive organs (pollen and seed cones) are not
oecious, evergreen. Resin in bark and phylloclades. always present but have some diagnostic characters
Bark smooth, becoming scaly. Branches in pseudo- necessary for correct determination. As a further
whorls at intervals along a main or single stem, pla- aide therefore the country or region of origin is also
giotropic and/or ascending (Rauhs model); lateral given with each species.
branches of highest order transformed to simple or
compound (pinnate) phylloclades (green planated 1a. Phylloclades simple or pinnately compound;
branchlets of variable size and shape) arranged spi- margins crenately or obtusely lobed. 2
rally or in pseudo-whorls. Indeterminate shoots 1b. Phylloclades pinnately compound; margins
ending in a globose bud with imbricate scales. True entire to deeply dissected 3
leaves on seedlings on the main stem, linear, chang- 2a. Phylloclades mostly simple. Pollen cones on a
ing gradually to subulate and scaly, inconspicuous short branching system, solitary or with 25
and deciduous on lateral branchlets in older plants. together. Native of Tasmania P. aspleniifolius
Stomata in mature plants restricted to phylloclades. 2b. Phylloclades pinnately compound or simple.
Pollen cones crowded in a low spiral at or below apex Pollen cones from a terminal bud with up to 10
of leading normal shoots, becoming more or less in a pseudo-whorl. Native of New Zealand
remotely placed with shoot elongation, subtended P. trichomanoides
by perular scales, stalked, cylindrical; microsporo- 3a. Seed cones 48(10) alternately on either side
phylls with a small triangular head and two rela- of the petiolate phylloclade base. Pollen cones
tively large pollen sacs containing bisaccate pollen. from a terminal bud with 1520 in a pseudo-
Seed cones (sub)marginal or terminal on petiolate whorl. Native of New Zealand P. toatoa
or foliate parts of phylloclades, in rows, in pairs or 3b. Seed cones on the margins or in an apical notch
solitary; consisting of a few to many bract scales, of of phylloclades, solitary or with 23 together.
which 1several are fertile; ovules axillary to a bract Pollen cones on a short branching system, up to
scale, solitary, erect, subtended by an aril, which 15 together. Native throughout Malesia
arises after the ovule has been formed. Seeds ovoid, P. hypophyllus
dorsiventrally compressed, partly embedded by the
fused, swollen and succulent bract scales of the cone
Phyllocladus aspleniifolius (Labill.) Hook. f., or rusty brown; new phylloclades bright green or
London J. Bot. 4: 151. 1845. Podocarpus asple- tinged red; old phylloclades lustrous deep green or
niifolius Labill., Nov. Holl. Pl. Sp. 2: 71, t. 221. dark green above, pale green below, sometimes glau-
1806. Type: Australia: Tasmania, North Esk River, cous. Stomata numerous on the underside (abaxial)
Cataract Gorge, [Habitat in capite Van-Diemen], in irregular lines. Pollen cones on a short branch-
J. J. H. de Labillardire s.n. (lectotype FI). Fig. 184 ing system from a terminal bud, solitary or with up
to 5 cones together, with one or two small bracts
Etymology (foliola) at their base, cylindrical, 58mm long,
22.5mm wide, pink or reddish when immature,
The species epithet compares this species with the becoming yellow. Microsporophylls ovoid-triangu-
foliage of Asplenium ruta-muraria, a small European lar, with two basal, globose pollen sacs. Seed cones 561
fern common on rocks and ancient walls. axillary to reduced scale leaves on margins or at
base of reduced phylloclades, or on terminal short
Vernacular names branching systems without phylloclades, solitary or
with 24 together, each consisting of several bracts,
Celery-top pine 25 of which are fertile and merge to a red or pur-
plish structure 35mm long, which becomes slightly
Description swollen and pinkish red (drying leathery brown).
Seeds 25 per cone, 1 per fertile bract, each in a white
Shrubs or trees to 20m tall, in tall forest with a aril covering the lower two third of seed; distal part
clear bole to 10m or more, d.b.h. to 50cm. Bark to free, 5mm long, semi-ovoid (laterally flattened) with
20mm thick, with large lenticels; outer bark deeply a lateral ridge and a small protrusion at apex, green-
furrowed and scaly in old trees, exfoliating in small ish black to black.
to medium sized flakes, dark brown weathering dark
grey or blackish; inner bark close to wood red or Distribution
pink, slightly fibrous. Branches spreading or ascend-
ing, forming a narrow or wide pyramidal crown. Australia: Tasmania.
Foliage branches mostly straight, spreading at an TDWG codes: 50 TAS
angle of less than 90, robust , terete, smooth, newest
shoots tinged red, becoming green then light brown, Ecology
terminating in a short bud with spreading, triangular
to acicular scales. True leaves on seedlings 1015mm Phyllocladus aspleniifolius occurs in montane tem-
long, subulate-linear, 1mm wide, with a midvein perate rainforest up to the tree line from near sea
and stomata on abaxial (lower) side, acute; along level on the west coast of Tasmania to 1200m a.s.l.
new shoots and on margins of phylloclades filiform in the central highlands. The largest trees grow in
leaves appear in young plants, 13mm long, decidu- mixed forest with Eucalyptus spp. at lower eleva-
ous. Phylloclades axillary to reduced, deciduous, fili- tions. At higher altitude (above 700800m a.s.l.)
form scale leaves, mostly simple, bifacially flattened, it is found in open woodland with Eucalyptus coc-
leaf-like, (1.5)2.55(8)cm long, mostly cuneate or cifera, Nothofagus cunninghamii, N. gunnii, Richea
rhombic in outline, with crenately to obtusely lobed pandanifolia, R. scoparia, Athrotaxis cupressoides,
margins, narrowing to a cuneate or petiolate base, A. selaginoides, A. laxifolia, and various shrubs.
sometimes more or less pinnatifid; phylloclades on Towards the tree line it grows with Orites acicu-
old trees smallest and least dissected, on seedlings laris, O. revoluta, Tasmannia lanceolata, Podocarpus
often pinnatisect. Venation of phylloclades penni- lawrencei, Diselma archeri, Pherosphaera hookeri-
parallel, with a midvein from base to (near) apex ana, Nothofagus gunnii, Olearia spp., and heath-
and few to numerous veins running parallel to each like dwarf shrubs and alpine herbs. The substrate is
other from midvein or midvein line to margins acidic and derived from dolerite, granite, or quarzite,
under a narrow but sometimes widening angle (i.e. and is usually well drained, such as thin soil on
they curve outwards). Flushing phylloclades reddish boulder or scree slopes. Precipitation is abundant
through much of the year, with no marked periods then light brown, terminating in a short bud with
of drought. spreading, triangular to acicular scales. True leaves on
seedlings 510mm long, subulate-linear, 0.50.7mm
Conservation wide, with a midvein and stomata on abaxial (lower)
side, acute; at base of phylloclades and on their mar-
IUCN: LC gins filiform leaves appear in young plants, 25mm
long, deciduous. Phylloclades axillary to reduced,
Uses deciduous, filiform scale leaves, pinnately com-
pound, the pinnate segments bifacially flattened,
Celery-top pine varies from a medium size tree in leaf-like, extremely variable in size and shape, 113cm
562 the forests at middle altitudes to a shrub in the sub- long, 0.55cm wide, generally rhombic in out-
alpine zone. The wood of good size trees is straight line but also parabolic to lanceolate, with entire to
grained and dense, pale brown, and not dissimilar deeply dissected margins (crenate to laciniate), the
to yew (Taxus). It is used for construction, flooring, latter type mostly on seedlings to saplings, narrow-
ship masts, furniture, and cabinet work. This species ing to a petiolate base; phylloclades with seed cones
is rare in cultivation, limited to botanic gardens and smaller than sterile ones, irregularly dissected, with
arboreta and a few private gardens. few lobes, often cuneiform or deeply emarginate.
Venation of phylloclade segments penni-parallel,
with a midvein from base to (near) apex and few to
Phyllocladus hypophyllus Hook. f., Icon. Pl., n.s., 5: numerous veins running parallel to each other from
t. 889. 1852. Type: Malaysia: Sabah, Ranau District, the midvein or midvein line to margins under a nar-
Mt. Kinabalu N.P., H. Low s.n. (holotype K). Fig. row but sometimes widening angle (i.e. they curve
185, 186 outwards). Flushing phylloclades yellow-green, red,
or rusty brown; new phylloclades bright green or
Etymology tinged red or copper, sometimes glaucous; old phyl-
loclades lustrous deep green or dark green above,
The species epithet is composed of Greek hypo = pale green below, sometimes glaucous. Stomata
below or under and phyllus = leaved. numerous on the underside (abaxial) in irregular
lines. Pollen cones at base of a new shoot or from a
Vernacular names terminal large bud, usually on a branching system
with up to 15 cones, each cone on a 525mm long
Celery pine, Celery-top pine; bindang, pelayo peduncle subtended by a strap-like, scarious bract,
(Borneo-Sarawak); kayu karongan, rapak-rapak sometimes with a few reduced phylloclades and with
(Borneo-Kalimantan); kayu empire (Sulawesi); beja- two small bracts (foliola) at their base, cylindrical,
lin (Maluku); dalung (Pilippines). 1015mm long, 3mm wide, pink or reddish when
immature, becoming yellow. Microsporophylls
Description ovoid-triangular, often acuminate, with two basal,
globose pollen sacs. Seed cones axillary to reduced
Shrubs or trees to 40m tall, in tall forest with a clear scale leaves on margins of phylloclades or in an api-
bole to 20m or more, d.b.h. to 100cm, usually more cal notch, or on terminal short branching systems
slender. Bark to 25mm thick, with large lenticels; without phylloclades or with a few much reduced
outer bark becoming scaly, exfoliating in small to phylloclades, solitary or with 23 together, each
medium sized flakes, dark brown weathering dark consisting of several bracts, 13 of which are fer-
grey or blackish; inner bark close to wood red or tile and merge to a red or purplish cupulate struc-
pink, slightly fibrous. Branches spreading or ascend- ture 57mm long, which becomes slightly swollen
ing, forming a narrow or wide crown much depend- and bright red (drying leathery brown). Seeds soli-
ing on location. Foliage branches mostly straight, tary to each fertile bract, with a white or yellow
spreading at an angle of less than 90, robust, terete, aril covering the lower half of seed only, 57mm
smooth, newest shoots tinged red, becoming green long, semi-ovoid (laterally flattened) with a small
protrusion at apex, ripening to lustrous tan or chest- light construction, flooring, interior finish, joinery,
nut brown. cupboards, and to a limited extent for the making
of furniture. It has excellent properties for plywood
Distribution and veneer. More specialized uses are for laboratory
bench tops, foundry patterns and storage batteries.
Malesia: Borneo, Maluku [Moluccas], Philippines, The resin (copal) has been collected by tapping the
Sulawesi; Papuasia: New Guinea. trees. The bark is used for roofing in New Guinea
TDW|G codes: 42 BOR-BR BOR-KA BOR-SB and the phylloclades are used to make tea in Borneo.
BOR-SR MOL PHI SUL 43 NWG-IJ NWG-PN This species is in cultivation, but rare and mainly
limited to botanic gardens.
Ecology 563
Phyllocladus hypophyllus occurs in lower mon- Phyllocladus toatoa Molloy, New Zealand J. Bot. 34:
tane to subalpine evergreen rainforests at altitudes 290. 1996. Phyllocladus glaucus Kirk, Trans. New
between (310)600m and 3400(4000)m a.s.l. At Zealand Inst. 1: 149. 1869 [glauca], non Carrire
lower altitudes it grows as a canopy tree of consid- (1855). Type: New Zealand: North Island, Great
erable size with other conifers, e.g. Agathis sp. in Barrier Island, Kiwiriki, T. Kirk WELT 37785
kerangas on white sand derived from sandstone, or (holotype WELT).
in mixed forests with Podocarpaceae, Fagaceae and
Lauraceae as the dominant families of trees. It is Etymology
also found in high montane cloud forest or mossy
forest, which remains lower than 20m and is char- Toatoa is the Maori name for this species.
acterized by epiphytic growth of ferns and mosses.
Conifers, including P. hypophyllus, Dacrydium sp., Vernacular names
Dacrycarpus sp., and Podocarpus sp. may dominate,
or these forests are mixed with angiosperms. In New Celery-top pine; toatoa (Maori)
Guinea Nothofagus grandis is often the dominant
tree species, with Phyllocladus and podocarps mixed Description
in. At still higher altitudes the forest is dwarfed and
P. hypophyllus becomes shrubby, often growing on Small trees to 15m tall, d.b.h. to 60cm. Bark with
the edges of boggy grasslands (especially in New large lenticels; outer bark becoming fissured and
Guinea) or on rocky ridges. This species is found scaly, exfoliating in small to medium sized flakes, dark
on a variety of substrates, such as granite, serpen- brown weathering dark grey or blackish. Branches
tine, sandstone, peaty soils, and sometimes volcanic ascending, forming a narrow or wide pyramidal or
deposits or eroded limestone. rounded crown. Foliage branches mostly straight,
spreading at an angle of less than 90, robust, terete,
Conservation smooth, newest shoots green then light brown, ter-
minating in a short bud with free, triangular, acu-
IUCN: LC minate and dorsally keeled scales. True leaves on
seedlings 510mm long, linear, 0.50.7mm wide,
Uses acute, often interspersed with juvenile, deeply pin-
natifid, 35cm long phylloclades; at base of phyllo-
As a timber tree this species is of local importance clades and on their margins filiform leaves appear in
only due to its scarcity as a big forest tree. In Papua young plants, 25mm long, deciduous. Phylloclades
New Guinea, a ban on the export of its unsawn tim- axillary to reduced, deciduous, filiform scale leaves,
ber is in force in order to stimulate domestic process- pinnately compound, pinnate segments bifacially
ing. The wood is very similar to that of Podocarpus flattened, leaf-like, 1.56cm long, 14cm wide,
and straight grained, fine textured and easy to work, rhombic to obovate-flabellate in outline, with entire
but non-durable for outdoor purposes. It is used for to deeply dissected margins (crenate to laciniate on
Uses Conservation
Distribution
Phyllocladus trichomanoides D. Don var.
trichomanoides. Type not designated. New Zealand: North Island, South Island.
TDWG codes: 51 NZN NZS
Description
Ecology
Trees to 23m tall. Phylloclades mostly pinnately
compound or pinnatifid. Pollen cones with 510mm Var. alpinus is found from ca. 500m a.s.l. upwards,
long peduncles. but in the far south and west of South Island it is
found at sea level as well.
Distribution
Conservation
New Zealand: North Island, N South Island.
TDWG codes: 51 NZN NZS IUCN: LC
Picea A. Dietr., Fl. Gegend Berlin 2: 794. 1824. Type: Picea abies (L.) H. Karst.
[Pinus abies L.] (Pinaceae).
Sect. Casicta Mayr 1b. Seed cones smaller, or if longer than 10cm less
(Type: P. jezoensis) than 5 cm wide (measured with opened seed
Subsect. Sitchenses E. Murray scales), with smaller seed scales 4
Species: P. sitchensis (type), 2a. Leaves very rigid, curved, 1.82.5 mm wide,
P. jezoensis, P. likiangensis, spreading radially or assurgent. Young shoots
P. linzhiensis, P. purpurea thick, very firm 3
Subsect. Pungentes E. Murray 2b. Leaves flexible, nearly straight, ca. 1mm wide,
Species: P. pungens (type), not assurgent. Young shoots thin and flexible
P. engelmannii, P. lutzii P. smithiana
3a. Vegetative buds large (812 48mm), ovoid
568 Key to the sections, subsections and series or ovoidoblong, smooth, shining chest-
of Picea nutbrown; leaves strongly curved and assur-
gent on thick branchlets P. torano
1a. Seed scales of cones thin and flexible, more or 3b. Vegetative buds smaller (56 3.54mm), con-
less papery, sometimes coriaceous, usually ical or ovoidconical, not smooth and shining,
undulate, with erose margins, loosely imbricate slightly pubescent; leaves not assurgent, on firm
before ripening of the cone Sect. Casicta 2 but less thick branchlets P. neoveitchii
1b. Seed scales of cones rigid, more or less thin 4a. Seed cones small (36 1.52.5cm), narrowly
woody, smooth or undulate, with entire or den- tapering at both ends, light redbrown when
ticulate margins, closely imbricate before rip- ripe; seed scales thin, convex, longer than wide,
ening of the cone Sect. Picea 3 with rounded or truncate apex; upper margin
2a. Leaves quadrangular in crosssection, nearly not erose. Leaves glaucous P. glauca
equifacial or bilateral, amphistomatic, glaucous 4b. Seed cones and seed scales different. Leaves
green or glaucous on all sides green or glaucous- green 5
Subsect. Pungentes 5a. Seed cones relatively small (not longer than
2b. Leaves more or less flattened, not equifacial, 8cm), with obtuse apex; seed scales small (max.
epistomatic or epiamphistomatic (with more 2 1.6cm), with usually rounded, entire upper
lines of stomata on the dorsal side of the leaf), margins 6
differently coloured on two sidesSubsect. 5b. Seed cones usually larger (but variable accord-
Sitchenses ing to altitude and latitude in several species!),
3a. Leaves epistomatic, usually distinctly flattened, seed scales variable 11
sometimes transversely rhombic in cross 6a. Leaves very dense above the shoot, curved 7
section Subsect. Omorikae 6b. Leaves more remote, leaving the shoot visible
3b. Leaves amphistomatic, quadrangular (equifa- from above, usually straight 8
cial) or rhombic (bilateral, keeled) in cross 7a. Leaves pressed forward above the shoot, the
section Subsect. Picea 4 apices curved towards it. Young shoots nearly
4a. Young shoots (densely) pubescent; leaves not white; buds small (34mm), slightly resinous
wider than 1(1.5)mm, not longer than 1.5cm. P. morrisonicola
Seed cones usually small Ser. Rubentes 7b. Leaves strongly assurgent, the apices curved
4b. Young shoots (mostly) glabrous; leaves wider upward. Young shoots yellowish to orange
than 1 mm (usually more than 1.5 mm), if ca. brown; buds larger, very resinous P. koyamae
1mm wide much longer than 1.5cm. Seed cones 8a. Young shoots (branchlets) pale buffgrey; buds
usually large Ser. Picea not resinous P. wilsonii
8b. Young shoots yellowish brown to orangebrown;
Key to the species (and some subspecies and buds usually slightly resinous 9
varieties, but excluding hybrids) of Picea 9a. Leaves small (813 11.4mm), spreading and/
Sect. Picea, Subsect. Picea, Ser. Picea or appressed forward above the shoot
P. maximowiczii
1a. Seed cones large, broad (815(18) 47 cm), 9b. Leaves larger, spreading radially 10
with very large, convex seed scales (23 10a. Young shoots (pale) yellowish brown or pale
1.53cm), with incurved, rounded margins 2 redbrown; leaves with obtuse or acute apex
P. koraiensis 19b. Buds not resinous (or only slightly so); bud
10b. Young shoots bright orange or redbrown; scales appressed 20
leaves with acute apex P. obovata 20a. Bark grey, inner bark orange P. schrenkiana
11a. Seed scales with a narrowly elongated, often 20b. Bark orangebrown to redbrown
emarginate apex (in P. abies var. abies cones P. crassifolia
with weakly elongated or even obtuserounded 21a. Young shoots pale yellowish brown, entirely
seed scales occur, but these types are less glabrous; buds slightly resinous 22
common) 12 21b. Young shoots orange or orangebrown, usually
11b. Seed scales not elongated at apex, but obtuse, variously pubescent; buds resinous 23
rounded or truncate 17 22a. Leaves with 35 lines of stomata on each face;
12a. Seed cones small (47.5cm long); apex of seed upper margin of seed scales entire. Resin canals 569
scales entire, reflexed in leaves usually present P. martinezii
P. alcoquiana var. reflexa 22b. Leaves with 410 lines of stomata on each face;
12b. Seed cones usually longer than 6 cm; apex of upper margin of seed scales denticulate. Resin
seed scales emarginate, (slightly) incurved 13 canals in leaves absent P. chihuahuana
13a. Seed cones 610(12) cm long. Leaves dense, 23a. Young shoots sparsely pubescent; buds gla-
curved forward; stomata in 13 lines on each brous at base. Seed scales not spreading wide or
ventral face, in 36 lines on each dorsal face reflexed in ripe cones 24
P. alcoquiana var. alcoquiana 23b. Young shoots often ferruginous pubescent;
13b. Seed cones 1216 cm long. Leaves radially buds pubescent at base. Seed scales spreading
spreading; lines of stomata equally numerous wide or slightly reflexed in ripe cones
on all sides of leaves 14 P. retroflexa
14a. Seed scales rhombicoblong; apex strongly 24a. Seed scales large (22.4 1.51.8 cm), with
elongated 15 nearly rounded apex P. aurantiaca
14b. Seed scales angularobovate (obtrullate) to 24b. Seed scales smaller (1.22 0.81.6 cm), with
more or less rhomboid; apex slightly elongated obtuse or rounded apex
16 P. asperata var. asperata
15a. Leaves (especially young leaves) glaucous
green; branches not pendulous Key to the species (and some subspecies and
P. asperata var. notabilis varieties, but excluding hybrids) of Picea
15b. Leaves (dark) green; branches spreading or Sect. Picea, Subsect. Picea, Ser. Rubentes
pendulous P. abies var. acuminata
16a. Young leaves glaucous green 1a. Seed cones narrowly cylindrical, tapering
P. asperata var. ponderosa towards apex, usually 59cm long. Leaves less
16b. Young leaves green P. abies var. abies than 1cm long, dark glossy green on all sides,
17a. Seed cones narrowly tapering towards apex; some white lines of stomata on ventral side,
seed scales rounded or truncate at apex. Young more lines
shoots pubescent P. alcoquiana var. acicularis on the dorsal side. Basal scales of buds obtuse-
17b. Seed cones obtuse at apex, more cylindrical; triangular P. orientalis
seed scales obtuse or rounded at apex. Young 1b. Seed cones ovoid, ovoidoblong or broad
shoots glabrous or pubescent 18 cylindrical, with obtuse or truncate apex.
18a. Seed cones narrowly cylindrical, usually 711 Leaves longer than 1 cm (if shorter, cones
2.53.5 cm, of very regular shape; seed scales smaller than
obovate, broad, with very regularly rounded 4 2.8cm), with a different colour. Basal scales
apex of buds acute-cuspidate 2
19 2a. Seed scales undulate, with undulate or emar-
18b. Seed cones broadly cylindrical, 814 35cm, ginate, erosedenticulate upper margin; cones
of less regular shape; seed scales with more or 3.58.5cm long, often with a truncate apex
less rounded or obtuse apex 21 P. glehnii
19a. Buds resinous; bud scales recurved near bud 2b. Seed scales not undulate, abaxial surface rough
apex P. meyeri or smooth, upper margin not emarginate; cones
smaller (max. 6cm long), with an obtuse apex 1b. Immature seed cones red or purple, ripening to
3 purplish brown or (dark) redbrown 3
3a. Seed cones very small (1.54 1.52.8 cm), 2a. Leaves thin (ca. 1mm), nearly quadrangular in
often numerous in the top of the tree, persisting crosssection, acute and pungent. Bracts
several years. Leaves usually 0.81.2 cm long, 58mm long (nearly half length of seed scales)
dark green or glaucous green aboveP. mariana P. sitchensis
3b. Seed cones usually larger (2.56 1.83.5cm), 2b. Leaves broader (1.52 mm), obtriangular in
soon deciduous after shedding seeds. Leaves crosssection, acutish or mucronate. Bracts
usually 11.5cm long, shiny light green above 45mm long (less than a third length of seed
P. rubens scales) P. jezoensis subsp. jezoensis
570 3a. Seed cones small (2.55 1.73 cm), purple,
Key to the species (and some subspecies and violet or crimson when immature; seed scales
varieties, but excluding hybrids) of Picea rhombic, with incurved, papery apex P.
Sect. Picea, Subsect. Omorikae purpurea
3b. Seed cones larger or with different seed scales4
1a. Leaves transversely rhombic in crosssection, 4a. Young shoots glabrous, reddish brown in the
acute and pungent at apex, up to 3.5cm long, second year; buds not resinous; leaves distinctly
radially spreading P. spinulosa flattened, epistomatic. Seed cones with nearly
1b. Leaves distinctly flattened, obtuse or acutish, obovate, flat seed scales
not pungent, pressed against the shoot above, P. jezoensis subsp. hondoensis
pectinate below 2 4b. Young shoots lighter, yellowish, or pubescent;
2a. Seed cones small (58 23 cm); seed scales buds (slightly) resinous; leaves variable. Seed
suborbicular, remaining imbricate. Young cones mostly with rhombic or less often with
shoots distinctly pubescent P. omorika angularobovate (obtrullate) seed scales 5
2b. Seed cones larger; seed scales of different shape, 5a. Young shoots brown pubescent, hairs glandu-
spreading after ripening of the cone. Young lar; leaves (almost) epistomaticP. linzhiensis
shoots glabrous to pubescent 3 5b. Young shoots glabrous or slightly pubescent,
3a. Seed scales obovate, convex, with a rounded, hairs not glandular; leaves amphistomatic
more or less incurved upper margin 4 P. likiangensis
3b. Seed scales obovateoblong or rhombic, with
(usually) recurved and sometimes elongated Key to the species (and some subspecies and
upper marginP. brachytyla varieties, but excluding hybrids) of Picea
4a. Leaves long and wide (1535 1.52 mm), Sect. Casicta, Subsect. Pungentes
curved and spreading. Branches of second
order extremely long pendulous. Seed cones 1a. Seed cones of medium size (58 34.5 cm);
sessile; seed scales regular, with rounded mar- seed scales obtrullate or rhombic; apex usually
gins and much resin P. breweriana narrowly elongated. Leaves more or less assur-
4b. Leaves smaller (1525 11.1 mm), straight. gent, rigid; apex very acute, pungent P.
Branches shorter and less pendulous. Seed pungens
cones with an oblique peduncle; seed scales 1b. Seed cones usually small (36 22.5cm); seed
obovate, but more irregularly shaped P. farreri scales obovateobtrullate; apex not narrowly
elongated. Leaves directed forward, flexible;
Key to the species (and some subspecies and apex acute but not pungent 2
varieties, but excluding hybrids) of Picea 2a. Leaves 1.52mm wide
Sect. Casicta, Subsect. Sitchenses P. engelmannii subsp. engelmannii
2b. Leaves 11.2mm wide
1a. Immature seed cones yellowish green or green, P. engelmannii subsp. mexicana
ripening to yellowish brown or (pale) reddish
rown 2
Picea abies (L.) H. Karst., Deutsche Fl.: 324. 1881. Taxonomic notes
expansion into W Europe. In the Alps Picea abies (Brgger) Stein, Gartenfl. 37: 346. 1887; Picea abies
occupies the montane to subalpine zones (depen- (L.) H. Karst. var. alpestris (Brgger) P. A. Schmidt,
dent on local climate) especially on moist sites and Haussknechtia 4: 38. 1988.
in cold air pockets. Although it can occur on most
substrates, acidic soils are most common and wide- Description
spread as is testified by the undergrowth, if present,
of ericaceous shrubs and sub-shrubs. Commonly Seed cones variable in size, (2.5)615(20)cm long;
growing with Picea abies in the boreal forests are seed scales ovoid-oblong to rhomboid-oblong; apex
Betula sp. and Populus tremula, with willows (Salix) variable but not narrowly elongated.
alongside streams and lakes. In the Alps Picea abies
572 occurs with Larix decidua, Pinus cembra, and P. syl- Distribution
vestris or P. nigra, if not in pure stands. In E Europe,
Picea abies is a constituent of mixed conifer-broad- Central, N and E Europe, eastward to the Ural
leaved woodland from the Bialowiecza Forest in Mountains.
the north to the valleys of the eastern Alps and the TDWG codes: 10 FIN NOR SWE 11 AUT-AU AUT-LI
Carpatians. CZE-CZ CZE-SL GER HUN POL SWI 12 FRA-FR 13
ALB BUL GRC ITA-IT ROM YUG-BH YUG-CR YUG-
Uses KO YUG-MA YUG-MN YUG-SE YUG-SL 14 BLR BLT-
ES BLT-KA BLT-LA BLT-LI RUC RUE RUN RUW
Norway spruce is an important timber tree in UKR-MO UKR-UK
Europe, where outside the boreal forest zone most
commercial timber is now harvested from planta- Conservation
tions or from managed forests in which other trees
are suppressed. Forestry has expanded the range of IUCN: LC
this species considerably further west. The wood is
used for pulpwood as well as construction, furni-
ture (most of the popular pine furniture is made Picea abies (L.) H. Karst. var. acuminata (Beck)
with wood from Norway spruce), and special uses Dallim. & A. B. Jacks., Handb. Conif., ed. 2: 390.
like the sound boards of pianos and the bodies of 1931. Picea excelsa (Lam.) Link var. acuminata
guitars and violins. The famous Stradivarius vio- Beck, Ann. K. K. Naturhist. Hofmus. 2: 61. 1887.
lins were made with wood of Norway spruce from Type: Illustration in M. Kienitz, ber Formen und
the Alps. In Europe this species is the most popu- Abarten heimischer Walbume, t. 3, f. 5a. 1879
lar Christmas tree, a tradition that actually started (holotype).
in Germany, with the extensive afforestation begin-
ning in the 18th century. Norway spruce is not much Description
planted as an amenity tree, but in horticulture more
than 200 cultivars have been selected, with differ- Seed cones generally large, 1216cm long; seed
ent habits including weeping, prostrate and dwarf scales rhombic, with narrowly elongated, acuminate
forms, red, white or yellow flushing leaf forms, and and incurved, erose-denticulate apex.
(other)monstrosities.
Distribution
2 varieties are recognized:
Europe: Jura, Alps, Carpathian Mts., S Norway,
Sweden?
Picea abies (L.) H. Karst. var. abies. Pinus abies TDWG codes: 10 NOR 11 AUT-AU CZE-CZ CZE-SL
L., Sp. Pl. 2: 1002. 1753. Type: Illustration Picea in POL ROM SWI 12 FRA-FR 13 ITA-IT 14 UKR-UK
Camerarius, Pl. Epit.: 47. 1586 (lectotype).
Conservation
Abies alpestris Brgger, Jahresber. Naturf. Ges.
Graubndens, ser. 2, 29: 167. 1886; Picea alpestris IUCN: LC
Distribution Description
574 Japan: central Honshu. New shoots pubescent; leaves curved forward, 0.8
TDWG codes: 38 JAP-HN 1.3cm long. Seed cones 47.5cm long; seed scales
entire, apically narrowed and reflexed.
Conservation
Distribution
IUCN: LC
Japan: Honshu (Akaishi Range).
TDWG codes: 38 JAP-HN
Picea alcoquiana (Veitch ex Lindl.) Carrire var.
acicularis (Maxim. ex Beissn.) Fitschen, in Beissner, Conservation
Handb. Nadelholzk., ed. 3: 258. 1930. Picea acicu-
laris Maxim. ex Beissn., Handb. Nadelholzk.: 380. IUCN: EN [B2ab (ii, iii, v)]
1891; Picea bicolor (Maxim.) Mayr var. acicularis
(Maxim. ex Beissn.) Shiras., Bot. Mag. (Tokyo) 27:
130. 1913. Type: Japan, coll. C. J. Maximowicz Picea asperata Mast., J. Linn. Soc., Bot. 37: 419.
(type not designated). 1906.
575
Plate 23. Picea alcoquiana. 1. Habit of tree. 2. Branch with foliage. 3. Seed cone (var. alcoquiana). 4. Seed
scale (var. alcoquiana). 5. Seed cone (var. acicularis). 6. Seed scale (var. acicularis). 7. Seed cone (var. reflexa).
8. Leaf. 9. Seeds.
recurved. Vegetative buds ovoid-conical or conical, TDWG codes: 36 CHC-SC CHI-NX CHN-GS CHN-SA
acute, 612 58mm, resinous; bud scales triangular CHQ
acute, keeled, appressed, or the apical scales slightly
recurved, yellowish brown with reddish margins, Ecology
persisting several years. Leaves spreading radially,
forward or assurgent above, parted and spreading Picea asperata occurs in the high mountains of W
laterally below, 12(2.5)cm long, 11.8mm wide central China, at elevations between 1500m and
(widest at base), linear, curved or straight, quad- 3800m a.s.l., usually above 2400m in Sichuan. The
rangular-rhomboid in cross-section, acuminate and soils are grey-brown mountain podzols. The climate
very pungent at apex; amphistomatic, with 34 lines is continental, subalpine, with cold winters and dry
576 of stomata on each face; colour of young leaves glau- summers (annual precipitation less than 500mm).
cous green, later dull green. Pollen cones near end It forms mostly pure forests on N-facing slopes, or
of shoots, axillary, 11.5cm long, reddish, then yel- mixtures with other species of Picea, in the south
low. Seed cones terminal, erect at first, pendulous of Gansu it may be mixed with Abies nephrolepis.
at maturity, short pedunculate or sessile, cylindri- Betula albo-sinensis is the most common broad-
cal, tapering at both ends (fusiform), obtuse at apex, leaved associate.
(5)612(15) cm long, (2.5)34(4.5) cm wide
with opened scales; colour (immature) purplish Uses
green or purple, ripening to (dark) brown or red-
dish brown. Seed scales obovate or rhombic-oblong, Picea asperata is an important timber tree in China.
1.22 0.81.6cm at mid-cone; surface striated The wood is mainly used for pulpwood and to a
to nearly smooth, glabrous; upper margin obtuse, lesser extent for construction. Old growth stands
rounded or truncate to emarginate, usually slightly of this potentially large spruce have been reduced
incurved; base cuneate. Bracts rudimentary, ligulate, to less accessible mountain slopes and valleys and
23mm, entirely included. Seeds ovoid, with acute plantation forestry has begun to replace the natural
apex, 24 1.52.8mm, dark brown or grey-brown; stands as a resource for spruce timber, but as yet on a
seed wings ovate-oblong, 812 56mm, yellowish scale that is incapable of meeting growing demands
brown, transparent. in Chinas rapidly growing economy. The species and
its varieties are in cultivation as amenity trees and in
Taxonomic notes arboreta and pineta in Europe and the USA, mostly
still based on seed collections by Ernest Wilson,
Picea asperata is a variable Chinese spruce, not Joseph Rock and some other early 20th century plant
unlike Picea abies in Europe, and it might be bet- hunters who travelled widely in southern Gansu and
ter taxonomic practice to recognize this as such and western Sichuan.
not to name varieties as they seem to appear in the
limited numbers of specimens (both in herbaria and 3 varieties are recognized:
grown from seed in arboreta) available for study.
Population based studies in the field on P. abies
have revealed regional forms or races, some related Picea asperata Mast. var. asperata. Type: China:
to environmental factors, as well as much intro- Sichuan, [Western China], E. H. Wilson E. H.
gression. If such detailed studies were available for 3025 (holotype BM).
P. asperata, similar patterns would be likely. As some
of the varieties have been traditionally recognized Description
and since the above suggested studies are not avail-
able, I have tentatively retained them here. New shoots glabrous or variously pubescent; bud
scales mostly appressed. Seed cones (5)612cm
Distribution long, (2.5)34cm wide when opened; seed scales
obovate-oblong, with obtuse, rounded or truncate
China: Gansu, Ningxia (Helan Shan), E Qinghai, SW upper margin.
Shaanxi, Sichuan.
Distribution Distribution
China: Gansu, Ningxia (Helan Shan), E Qinghai, China: W Sichuan (Panlong Shan).
SW Shaanxi, Sichuan. TDWG codes: 36 CHC-SC
TDWG codes: 36 CHC-SC CHI-NX CHN-GS CHN-SA
CHQ Conservation
Young shoots glabrous, bright orange brown; bud Orange spruce; baipi yun shan (Chinese)
scales loosely imbricate, apically reflexed, yellowish
brown. Seed scales rhombic oblong, emarginate. Description
a cute-mucronate, pungent; amphistomatic, stomata in the lower elevations of the SE of its range to only
in 4 bands of 36 lines; young leaves bluish green, 500700mm in the NW. It occurs in mixed conif-
later grey-green. Pollen cones near end of shoot, erous forest, with e.g. Picea likiangensis var. rubes-
axillary, 1.52cm long, reddish, then yellow with cens, Abies squamata, A. chensiensis, A. recurvata,
pollen. Seed cones terminal, erect at first, pendulous Tsuga chinensis, and locally, Larix potaninii. Betula
at maturity, sessile or short pedunculate, cylindrical- spp. are the common broad-leaved trees, while Pinus
oblong, obtuse at apex, 1012cm long, 44.5cm spp. occur mostly after disturbances and at the lower
wide with opened scales; immature cones bright elevations.
red, maturing to reddish brown; ripe cones shining
chestnut brown or dull brown. Seed scales rhombic- Conservation
578 ovoid, with nearly rounded apex, 22.4 1.51.8cm
at mid-cone; surface striated, shining, glabrous; The limited areas of occurrence and occupancy
upper margin slightly erose denticulate; base cune- (EOO and AOO) estimated for this species and the
ate. Bracts rudimentary, ligulate, 24mm, entirely inferred decline from logging operations and exten-
included. Seeds ovoid-conical, 34mm long, brown; sive deforestation in the area justify the Red List cat-
seed wings ovate oblong, 1214 56mm, transpar- egory Endangered.
ent, yellowish brown. IUCN: EN (A2cd)
Masters (op. cit.), in his lecture to the Linnean A timber tree of which no further details of its uses
Society, separated this species from the closely are recorded, presumably because it is not recog-
related species P. asperata Mast. and P. retroflexa nized as distinct from Picea asperata. It must be
Mast., all three newly described there. He already logged with this and other spruces and put to the
made allegations about their close relationship and same uses. It was introduced in England and is still
possible intermediate forms. These taxa have been found in some arboreta in the south, growing par-
variously treated as independent species or variet- ticularly well on shallow soil over chalk.
ies of P. asperata (Schmidt-Vogt, 1977). In Flora of
China 4: 28 (1999) and in Higher Plants of China 3:
37 (Fu et al., 2000) P. aurantiaca is treated as a vari- Picea brachytyla (Franch.) E. Pritz., Bot. Jahrb.
ety of P. asperata, while other accounts (e.g. Fu & Jin, Syst. 29: 216. 1900.
1992; Farjon, 1990, 1998, [2001]) have maintained the
species rank. A re-examination of relevant collec- Etymology
tions and populations seems desirable; this should
include work on DNA sequences, looking for mark- The species epithet (Greek: brachy = short; tylo =
ers which may help to distinguish species. with lumps or projections) refers to the short pul-
vini on the shoots.
Distribution
Vernacular names
China: W Sichuan (Zheduo Shan W of Kangding,
from Simaqiao to Xinyulingong). Sargents spruce; mai diao yun shan (Chinese)
TDWG codes: 36 CHC-SC
Description
Ecology
Trees to 30(40)m tall, d.b.h. to 11.2m; trunk
Picea aurantiaca is a subalpine species, occur- monopodial, straight; bark becoming rough, scaly,
ring between 2600m and 3800m a.s.l. (4000m flaking, dark grey, with reddish brown freshly
according to Rehder & Wilson, 1914). It is mostly exposed bark plates. Branches of first order long,
found on calcareous soils. The climate is cold and slender, spreading horizontally; branches of sec-
precipitation varies from high (no figures recorded) ond order slender, pendulous; crown broad conical,
open in old trees. Branchlets slender, flexible, lead- influence is stronger. It is a constituent of the montane
ing shoots stout, creamy white at first, later light coniferous forest of the eastern parts of the Himalaya
brown to orange-brown, sometimes pruinose; sur- and the mountains of the SW Plateau of China
face finely ridged and grooved, glabrous or minutely (Wang, 1961), with Abies densa, A. forrestii, Picea
pubescent; pulvini small, whitish yellow, at 4560 likiangensis, Pinus wallichiana, Tsuga dumosa, and
to the shoot axis. Vegetative buds ovoid conical, Larix potaninii as major species. Taxus wallichiana
on leading shoots 58 4 6mm, smaller on lateral is commonly found as an understorey tree in the
shoots, covered by leaves, (slightly) resinous; bud Himalayan part of its range.
scales closely appressed, triangular, chestnut brown,
persistent. Leaves curved forward and downward, Uses
with lower leaves parted, nearly pectinate, on con- 579
ing shoots more radially spreading, (0.8)12(2.4)cm This species is a timber tree, used for construction,
long, 11.5(2)mm wide, truncate at base, linear, interior flooring, aircraft, machines, furniture, and
curved, slightly flattened, keeled on both sides; apex wood pulp for the paper industry. In China this spe-
acute or mucronate; stomata on lower side only, in cies has been intensively exploited, depleting the
2 narrow bands separated by a midrib; leaf colour natural stands, and now is cultivated for afforesta-
dark green above, white bands below. Pollen cones tion. In Europe and North America it is often pres-
axillary, 12cm long, yellowish when shedding pol- ent in larger arboreta, mainly derived from seed
len. Seed cones terminal, erect at first, pendulous collections made by European plant hunters who
at maturity, short pedunculate, cylindrical-oblong traveled in China in the early decades of the 20th
or ovoid-oblong, (obliquely) tapering at base; apex century. It is a more attractive and shapely species
obtuse, (5)610(15)cm long, 34(5)cm wide than most spruces and new introductions from dif-
with opened scales colour (immature) green or ferent parts of its range should be recommended for
purplish green, ripening to dark brown with a pur- horticultural uses; it is extremely hardy and tolerant
plish band across each scale. Seed scales angular- of poor soils.
bovate to nearly rhombic, thin, rigid, 1.62(2.5)
11.4(1.8)cm at mid-cone; abaxial surface stri- 2 varieties are recognized:
ated or wrinkled, glabrous; upper margin variably
undulate or rounded to truncate, usually recurved; Picea brachytyla (Franch.) E. Pritz. var. brachytyla.
apex emarginate or erose; base cuneate. Bracts rudi- Abies brachytyla Franch., J. Bot. (Morot) 13 (8): 258.
mentary, ligulate, 23mm, entirely included. Seeds 1899. Types: China, Sichuan, P. Farges 806; Yunnan,
ovoid-oblong, 3 2mm, light brown; seed wings A. Delavay 4129 (syntypes P).
ovate-oblong, 1014 57mm, orange-brown.
Picea pachyclada Patschke, Bot. Jahrb. Syst. 48: 630.
Distribution 1913; Picea brachytyla (Franch.) E. Pritz. var. pachy-
clada (Patschke) Silba, Phytologia 68: 39. 1990.
China: S Gansu, NW Hubei, S Shaanxi, W Sichuan,
NW Yunnan, SE Xizang [Tibet]; NE India: Arunachal Description
Pradesh (Assam Himalaya); N Myanmar [Burma];
Bhutan (?). Seed cones 610(12)cm long, 34cm wide when
TDWG codes: 36 CHC-HU CHC-SC CHC-YN opened; seed scales angular-obovate, 1.62cm long,
CHN-GS CHN-SA CHT 40 EHM-AP 41 MYA 11.4cm wide; upper margin variably undulate,
emarginate or erose, usually recurved.
Ecology
Distribution
Picea brachytyla is a high mountain species, occur-
ring between 1300m and 3800m a.s.l. The soils are China: Chongqing, S Gansu, NW Hubei, S Shaanxi,
grey-brown mountain podzols. The climate is cold Sichuan, NW Yunnan, SE Xizang [Tibet].
and wet, with annual precipitation from 1000mm TDWG codes: 36 CHC-CQ CHC-HU CHC-SC
(N) to more than 2500mm (S), where the monsoon CHC-YN CHN-GS CHN-SA CHT
Picea brachytyla (Franch.) E. Pritz. var. complanata Picea breweriana S. Watson, Proc. Amer. Acad.
(Mast.) W. C. Cheng ex Rehd., Man. Cult. Trees, ed. Arts 20: 378. 1885. Type not designated.
2: 30. 1940. Picea complanata Mast., Gard. Chron.,
ser. 3, 39: 146. 1906. Type: China: Sichuan, Etymology
W Sichuan, E. H. Wilson 3031 (sheet No. 1, lecto-
580 type K, designated here). This species was named after William H. Brewer
(18281910) of the California State Geological survey.
Picea brachytyla (Franch.) E. Pritz. forma. rhom-
bisquamea Stapf, Bot. Mag. 148: sub t. 8969. 1923. Vernacular names
Picea ascendens Patschke, Bot. Jahrb. Syst. 48: 632.
1913; Picea brachytyla (Franch.) E. Pritz. var. ascen- Brewer spruce, Weeping spruce
dens (Patschke) Silba, J. Int. Conifer Preserv. Soc. 7
(1): 28. 2000. Description
wide with opened scales; colour (immature) green, Especially the smaller, isolated sub-populations
maturing to purplish brown, ripening to red-brown would be at risk.
or dull brown, very resinous. Seed scales obovate- IUCN: VU [B2b (ii, iii, v)]
flabellate, convex, thin but rigid, opening very wide
when dry, 1.52 1.31.8cm at mid-cone; surface of Uses
abaxial side smooth, slightly striated in some cones,
glabrous; upper margin entire, rounded or truncate, Brewer spruce is of little or no value as a timber tree.
slightly incurved; base cuneate. Bracts rudimentary, It may occasionally be logged with other conifers and
ligulate, 3mm, entirely included. Seeds ovoid, 34 put to use as pulp wood. In horticulture, however, it
23mm, brown; seed wings ovate, 79 56mm, is considered a very desirable ornamental tree due
orange-brown. to its long pendulous foliage branches. It is widely 581
planted especially in European arboreta and parks,
Distribution and generally performs well, although trees grown
from seed are slow to mature to the characteristic
USA: NW California, SW Oregon (Siskiyou Mts., habit; grafted trees develop faster. Few cultivars have
Klamath Mts.). been selected.
TDWG codes: 73 ORE 76 CAL
Qinghai spruce; Qinghai yun shan (Chinese) China: Gansu, Qinghai (Qilian Shan, around
Qinghai Hu), Nei Mongol (Daqing Shan), Ningxia
Description (Helan Shan).
TDWG codes: 36 CHI-NM CHI-NX CHN-GS CHQ
Trees to 2025m tall, d.b.h. to 5060cm; trunk
monopodial, straight; bark on trunk rough and scaly, Ecology
red brown. Branches of first order short, spread-
ing horizontally or upturned; branches of second Picea crassifolia occurs in high mountain ranges
order short, firm, spreading or ascending; crown of Central Asia, mainly on N-facing slopes, above 583
pyramidal or (narrowly) conical, open in old trees. steppe or desert, at elevations between 1600m and
Branchlets short, firm, leading shoots thick, pale 3800m a.s.l. It occurs on calcareous and non-calcar-
orange-yellow or greenish yellow, often pruinose, eous soils (the first soil type e.g. in Helan Shan). The
later grey, with prominent ridges and deep grooves, climate is cold continental and dry, with most of the
glabrous or with some scattered pubescence; pulvini precipitation falling as snow. It forms mostly pure
large, 22.5mm, erect or recurved. Vegetative buds forests, here and there with Betula albosinensis and
ovoid-globose, more conical on leading shoots, up groves of Populus tremula.
to 812 610mm, not or slightly resinous; bud
scales triangular, acutish, keeled, with erose mar- Conservation
gins, orange brown, often pruinose, persistent, leav-
ing wide collars of perular scales at base of shoots. IUCN: LC
Leaves crowded above shoot, upper leaves directed
forward and lower leaves curved upward, very rigid, Uses
(0.9)1.22.2(2.5)cm long, 1.52.5(3)mm wide,
linear, curved or nearly straight, nearly quadrate- Qinghai spruce is probably only locally exploited for
rhombic in cross-section, keeled on two sides; its timber (firewood?), as it occurs remote from any
apex obtuse-acutish; amphistomatic, stomata in 4 major urban centres and even major roads. It is not
bands; leaf colour bright green, with two whitish uncommon in cultivation in botanic gardens and
green stomatal bands. Pollen cones axillary, 11.5cm arboreta in Beijing, St. Petersburg and Moscow. In
long, yellowish pink. Seed cones terminal, erect at the West it is now also appearing, but still very rare;
first, pendulous at maturity, sessile, ovoid-oblong some trees are possibly misidentified, e.g. as Picea
or cylindrical, obtuse at apex, (5)711cm long, asperata.
2.53.5cm wide with opened scales; colour (imma-
ture) purplish red, maturing to green, with purple
margins of seed scales, ripe cones light brown or Picea engelmannii Parry ex Engelm., Trans. Acad.
dark brown. Seed scales broadly obovate-flabellate, Sci. St. Louis 2: 212. 1863.
slightly convex, flattened when open, 1.52
11.7cm at mid-cone; surface smooth, usually finely Etymology
striate, or slightly wrinkled, glabrous; upper mar-
gin entire, slightly incurved, rounded; base cune- W. E. Parry named this species after the botanist
ate. Bracts rudimentary, ligulate, 2mm, entirely George Engelmann (18091884), but failed to pro-
included. Seeds ovoid-oblong, 33.5 mm long, vide a validating description, an omission made
brown; seed wings ovate-oblong, 1013 45mm, good by Engelmann.
orange-brown.
Vernacular names
Engelmann spruce
Description Ecology
Trees to 4045(50)m tall, d.b.h. to 11.5(2)m; Picea engelmannii is widespread in the Rocky
trunk monopodial, straight; bark reddish brown, Mountains, from 600m to 3700(4000)m a.s.l.,
later grey with light brown and becoming rough the upper limit progressively higher from N to S. It
and fissured. Branches of first order short, slender, grows on various mountain soils, both calcareous
spreading horizontally, curved upward at ends, lower and non-calcareous. The climate is cold and humid
branches more pendant; branches of second order (precipitation above 600mm annually), with long,
short, dense, spreading or pendulous; crown nar- snowy winters and short, cool summers. The species
rowly conical or narrowly columnar, especially in the forms extensive pure forests or mixed coniferous for-
584 N or at high elevations, with branches to the ground ests, with Abies lasiocarpa, Pinus spp., Pseudotsuga
or the bole free of branches for a third to half in dense menziesii, Larix occidentalis, or Picea glauca as most
forest stands. Branchlets slender, firm, becoming common associated species.
pendulous, greenish yellow at first, soon yellowish
brown, ridged and grooved, finely pubescent when Uses
young; pulvini well developed, 2mm long, standing
at nearly 90 to shoot axis. Vegetative buds ovoid- Engelmann spruce is an important timber tree with
conical, 56mm long, resinous at apex; bud scales a high yield potential especially in managed stands
obtuse-triangular, appressed, apices later spreading, within its native range. Its knotty wood is not of very
red brown, persisting several years. Leaves spread- high grade, but nevertheless increasingly used for
ing radially, crowded above leading shoots, directed home building, carpentry, furniture, plywood, and
forward, (1.2)1.52.5(3)cm long, (1)1.52mm specialist uses such as musical instruments (pianos,
wide, linear, straight or slightly curved, quadrate- violins). Uses for mining timber, railroad sleepers,
rhombic in cross-section; apex acute (not pungent); and telephone poles have declined and mass pro-
amphistomatic, 2 narrow bands of 23 lines above, 2 duction is now directed to the pulp wood industry,
bands of 46 lines below; leaf colour glaucous green. especially in western Canada. Here massive clear
Pollen cones axillary, 11.5cm long, yellowish. Seed cut operations still bare whole mountainsides regu-
cones terminal, erect at first, pendulous at matu- larly. This species is rarely planted as an ornamental,
rity, sessile, ovoid-cylindric; apex obtuse, (2.5)3 although it will grow well even on poor soils, and
6(7.5)cm long, 22.5(3.5)cm wide with opened only a few cultivars are known in the trade. Spruces
scales; colour (immature) green tinged with red, are not much in use as Christmas trees in North
maturing to light reddish brown or pale yellowish America, unlike in Europe where they are the com-
brown. Seed scales obovate-obtrullate, thin and flex- monest genus for this purpose. The subspecies mexi-
ible, 1.21.5 0.91.2cm at mid-cone; abaxial surface cana is rare in cultivation.
smooth or finely striated, often undulate, glabrous;
upper margin rounded or truncate, undulate, entire 2 subspecies are recognized:
or erose-denticulate, sometimes lacerate; base cune-
ate. Bracts broadly ovate, cuspidate, 36mm long,
entirely included. Seeds ovoid, 23mm long, greyish Picea engelmannii Parry ex Engelm. subsp. engel-
brown; seed wings ovate-oblong, 1012 45mm, mannii. Picea glauca (Moench) Voss subsp. engel-
yellowish brown. mannii (Engelm.) T. M. C. Taylor, Madroo 15: 114.
1959; Picea glauca (Moench) Voss var. engelmannii
Distribution (Engelm.) Boivin, Naturaliste Canad. 93: 272. 1966.
Type: USA: Wyoming, Wind River Mountains,
W North America: Rocky Mountains from British F. V. Hayden s.n. (lectotype MO).
Columbia to North Mexico.
TDWG codes: 71 ABT BRC 73 COL IDA MNT ORE Picea engelmannii Parry ex Engelm. var. glabra
WAS WYO 76 ARI CAL NEV UTA 77 NWM TEX 79 Goodman, Madroo 10: 177. 1950.
MXE-CU MXE-NL
Description Distribution
Bark reddish brown, later grey with light brown. Mexico: S Chihuahua, Coahuila, Nuevo Len; USA:
Leaves 1.52mm wide. Bract scales of seed cones Arizona, Chiricahua Mts.
35mm long. TDWG codes: 76 ARI 79 MXE-CO MXE-CU MXE-NL
Distribution Conservation
W North America: Rocky Mountains from British The main population in the Sierra de la Marta (the
Columbia to New Mexico. type locality) was nearly exterminated in a forest fire
TDWG codes: 71 ABT BRC 73 76 77 in 1975. Other populations, e.g. on Cerro Mohinora 585
in Chihuahua, are much smaller and declining, and
Conservation even more susceptible to destructive fires. In addi-
tion, trees have been felled while regeneration is
IUCN: LC poor and slow.
IUCN: EN [A4acd; B2a (ii, iii, v)]
This taxon has first been found in the Sierra de Trees to 3035m tall, d.b.h. to 6070cm; trunk
la Marta, 75 km SE of Saltillo in NE Mexico, at monopodial, straight or slightly curved; bark on
3000m to 3400m a.s.l. on moist, N-facing, steep trunk rough and scaly, with small plates, grey or
slopes of dolomite limestone. Taylor & Patterson greyish brown. Branches of first order long, slender,
(1980) described a Picea hybrida population spreading horizontally or curved down, pendant
in S Chihuahua (Cerro Mohinora). D. S. Correll at the ends; branches of second order long, slen-
& H. S. Gentry 23183 (MEXU) from that loca- der, pendulous; crown broadly conical. Branchlets
tion (3100m) has cones (7cm) resembling large slender, drooping or pendulous, olive brown to pale
cones of P. engelmannii and leaves narrower than orange-brown, with flat ridges and shallow grooves,
1.3mm, with an acute (not pungent) apex. This sparsely pubescent at first, soon glabrous; pulvini
material is best treated as P. engelmannii subsp. weakly developed, 0.61mm long, at 4060 from
mexicana. shoot, pubescent, orange-brown. Vegetative buds
ovoid, or ovoid-conical, on leading shoots 45mm,
on lateral shoots smaller, slightly resinous; bud
scales obtuse, with erose margins, chestnut brown or Myanmar [Burma] leading up to a pass crossing into
purplish brown, persisting several years. Leaves on the deep valley of the Salween in Yunnan, China.
the upperside of shoots radially spreading forward, It was then also found on that side of the border,
more pectinate below, leaves subtending lateral so now two or perhaps three (sub)populations are
buds longest and at 90, (1.5)1.82.3(2.5)cm long, known to exist. Forest cutting and clearing for agri-
11.1mm wide, linear, straight or slightly curved, culture are expanding into higher elevations, threat-
convex above, flattened and keeled below, acute and ening the remaining montane coniferous forest
slightly pungent at apex; epistomatic, stomata in 2 and therewith this rare species. The status of the
bands of 56 lines on lower side; leaf colour green Burmese population(s) remains poorly known as
or glaucous green above, two niveous white stoma- no foreign botanists have visited this remote valley
586 tal bands below. Pollen cones axillary, 1.52.5 0.3 since the 1930s.
0.4cm, yellowish. Seed cones terminal, erect at first, IUCN: VU (D2)
pendulous at maturity; peduncles oblique, 5mm
long; shape ovoid-oblong to ellipsoid-cylindrical, Uses
obliquely tapering at base, obtuse at apex, (4)6
12(14)cm long, 2.54.5cm wide, colour (immature) Farrers spruce may be used locally for construc-
green or purplish green, maturing to dark brown. tion timber. Reginald Farrer introduced its seed to
Seed scales obovate-oblong or obovate, slightly con- England, where it was planted in several arboreta
vex, lateral margins recurved in opened cones, 1.2 and private parks. The resulting trees were believed
2.2 0.81.5cm at mid-cone; surface finely striated to have been lost until a tree at Exbury Gardens in
or undulate and wrinkled, glabrous; upper margin Hampshire, England was identified in 1979 by Chris
irregularly rounded, sometimes apically narrowed Page and Keith Rushforth as having grown from
and recurved, entire; base cuneate. Bracts rudimen- seed under Farrers collection number 1435, the same
tary, ligulate, 23mm long, entirely included. Seeds as the holotype specimen at the Herbarium of the
ovoid-oblong with narrowed apex, ca. 4 2.5mm, Royal Botanic Garden, Edinburgh (E). Herbarium
dark brown; seed wings ovate-oblong, 1315 6mm, specimens from this tree were used to illustrate
lustrous orange-brown. Picea farreri in my book Pinaceae (Farjon, 1990: 276)
and deposited in the Herbarium at the University of
Distribution Utrecht (U), now in the National Herbarium of the
Netherlands. This tree subsequently died, but cut-
China, W Yunnan (Salween River); Myanmar tings were taken, rooted or grafted, and distributed
[Burma] (Fen Shui Ling [valley & pass]). among arboreta and pineta in England and abroad.
TDWG codes: 41 MYA New plants were introduced in Scotland from
sources in Yunnan, China recently, but this species
Ecology remains extremely rare in cultivation.
Etymology Ecology
The nothospecies epithet refers to Finland (in a wider The dominant conifer in the northern taiga of
sense than the present territory of that country). European Russia, in drier sites mixed or occasion-
ally replaced by Pinus sylvestris, with undergrowth
Vernacular names of Vaccinium spp. or even lichens, in waterlogged
sites by Sphagnum bogs. Occurring from near sea
Karelian spruce level to 600800m a.s.l. depending on latitude,
dwarfed and becoming scattered at the Arctic tree
Description line. Large parts of its range are underlain by per-
mafrost; the deepest summer frost-free soils here 587
Trees; habit, bark and foliage similar to Picea abies are on river floodplains, where Picea attains its larg-
of northern Scandinavia. Seed cones symmetrical, est size. Frequently assosiated with Populus tremula
ovoid-oblong, 57.5cm long, 34cm wide when and Salix spp., especially after fires and slowly
opened; base flattened or tapered to a short pedun- replacing these pioneers in undisturbed sites, with
cle; seed scales spreading widely, obovate, convex; natural cycles of forest fires ranging from 40 years
apex more or less rounded to slightly elongated, to occasionally 400 years. On river floodplains
entire or erose, usually incurved. Alnus glutinosa characterizes a stadium between
pioneer vegetation and returning spruce forest,
Taxonomic notes especially in more sourthern parts of the range of
P. fennica.
The spruces that occur in a broad zone between
Norway spruce (Picea abies) and Siberian spruce (P. Conservation
obovata) have often been recognized as belonging
to the latter species (provided that it was accepted IUCN: NE
as distinct at the species rank from P. abies). The
extraordinary variability observed in the cone mor- Uses
phology of P. abies in Central and W Europe (see
e.g. Schmidt, 1991) appears to be absent in P. obovata This spruce is an important timber tree; in the Komi
in Siberia, as well as in the spruces of the transi- region of N Russia it is the mainstay of the economy
tion zone, the borders of which are poorly defined. in a thinly populated region. The production of tim-
However, certain character states, especially in the ber amounted to 6.8million m3 in the year 1940,
shape of the seed scales and their margins, are simi- a peak reached just before the Great Patriotic War
lar to P. abies, while the cones are mostly smaller, as (WW II) commenced and diverted all manpower
in P. obovata. The history of retreats of the forests to the fronts. Traditional houses there were all built
before the expansion of the Scandinavian ice cap with spruce logs and this use is making a come-
during ice ages, and their subsequent return in inter- back in all of northern Russia. Much of the tim-
glacials, can have caused an intermingling of these ber is processed to wood pulp for use in the paper
two closely related species in refugia, resulting in industry. Trees attributed to this hybrid species are
a hybrid taxon, as is borne out by isozyme studies in cultivation mainly around the Baltic Sea and in
(Schmidt, 2002a, b). Russia.
Distribution
Picea glauca (Moench) Voss, Mitt. Deutsch.
NE Russia: Archangelsk, Karelia, Kola Peninsula, Dendrol. Ges. 1907 (16): 93. 1907.
Komi; Finland?
TDWG codes: 14 RUN Etymology
Distribution Etymology
Boreal North America: from Newfoundland and This species commemorates Peter von Glehn
New York to NW. Alaska and W. Montana. (18351876).
TDWG codes: 70 ASK NUN NWT YUK 71 ABT BRC
MAN SAS 72 LAB NBR NFL-NE NFL-SP NSC ONT Vernacular names
PEI QUE 73 MNT 74 MIN SDA WIS 75 MAI MIC NWH
NWY Sakhalin spruce; aka-matsu (Japanese); el Glena
(Russian)
Conservation
Description
IUCN: LC
Trees to 30m tall, d.b.h. to 6070cm; trunk mono-
podial, straight; bark soon flaking, reddish brown,
Picea glauca (Moench) Voss var. albertiana (S. becoming rough and scaly, breaking into irregular
Br.) Sarg., Bot. Gaz. (Crawfordsville) 67: 208. 1919. plates, grey-brown to purplish grey or dark grey.
Picea albertiana S. Br., Torreya 7: 126. 1907; Picea Branches of first order long, slender, spreading
glauca (Moench) Voss subsp. albertiana (S. Br.) horizontally, but lower branches bent downward;
P. A. Schmidt, Haussknechtia 4: 38. 1988. Type not branches of second order dense, spreading horizon-
designated. tally; crown pyramidal or conical. Branchlets short,
slender, firm, orange or reddish brown, becoming
Description purplish brown; surface ridged and grooved, pubes-
cent in the grooves and on well developed, 1mm
New shoots pubescent; leaves 1.52cm long; apex long, densely set pulvini. Vegetative buds ovoid or
obtuse. Seed cones ovoid or ovoid-oblong, abruptly ovoid-conical, 46 4mm, (slightly) resinous; bud
narrowing at base. scales acute-triangular, basal scales long cuspidate,
red-brown, shiny, persisting several years. Leaves
radially spreading, the leaves above shoot directed
tal bands below. Pollen cones axillary, clustered on wood properties are similar to those of P. sitchensis,
pendulous shoots, 1.52cm long, yellow. Seed cones but this spruce does not attain the great dimen-
terminal, erect at first, pendant at maturity, often sions of that species. Much of its wood is processed
clustered, numerous, sessile or very short peduncu- to pulp for the paper industry, but more specialized
late, cylindrical, sometimes slightly curved, obtuse uses are furniture making and (in Japan)musical
at apex, (3)47(9)cm long, 23.5cm wide with instruments. The Ainu string instrument tonkori has
opened scales; immature cones green, ripening to a body made from Yezo Spruce. Log houses are con-
light or darker yellowish or reddish brown, some- structed with its wood in northern parts of its range,
times pruinose. Seed scales numerous, small, open- such as in Kamchatka where a small area of taiga is
ing wide, obovate-oblong to obtrullate or nearly dominated by this species. This spruce is commonly
rhombic, very thin and papery, but rigid, 11.2 0.6 used in afforestation especially in Japan and Korea; 591
0.8cm at mid-cone; surface finely striated, undulate in other parts of the Northern hemisphere it is less
or smooth and flat, glabrous; upper margin thin, often planted, either for forestry or for horticul-
undulate, erose-denticulate, flat or incurved; base ture. In western Europe, provenances from Honshu
cuneate. Bracts small, ligulate, with toothed and cus- are the only ones planted due to late frost damage
pidate apex, purplish, 45mm long, included. Seeds experienced with trees from more northern sources.
ovoid-cuneate, 3 2mm, light brown; seed wings Foresters have experimented with hybridization, e.g.
ovate-oblong, 610 45mm, light orange-brown with P. glauca and P. mariana and with its closest
or yellowish brown. relative, P. sitchensis. Only a few cultivars are known,
usually sporting yellow new foliage.
Distribution
2 subspecies and 2 varieties are recognized:
NE. China: coastal part of Jilin; Japan: Hokkaido,
Honshu; North Korea; Russian Far East.
TDWG codes: 31 KAM KHA KUR MAG PRM SAK 36 Picea jezoensis (Siebold & Zucc.) Carrire subsp.
CHM-JL 38 JAP-HK JAP-HN KOR-NK jezoensis var. jezoensis. Abies jezoensis Siebold &
Zucc., Fl. Japon. 2 (2): 19, t. 110. 1842. Type: Japan:
Ecology [Abies No. 2, Coniferae ex insula Jezo], P. F. von
Siebold s.n. (lectotype L).
Picea jezoensis occurs from near sea level to 2700m
a.s.l. (subsp. hondoensis on Honshu: 1100m to Picea ajanensis Fisch. ex Carrire, Trait Gn.
2700m), on various (podzolic) soils. The climate is Conif.: 259260. 1855; Picea jezoensis (Siebold &
cold temperate, moist or wet (1000mm to 2500mm Zucc.) Carrire var. ajanensis (Fisch. ex Carrire)
annual precipitation on Honshu). It is usually mixed W. C. Cheng & L. K. Fu, Fl. Reipubl. Pop. Sin. 7: 162.
with other conifers, e.g. Abies spp., Larix spp., Pinus 1978.
spp. (with P. pumila in the north of the range), and Picea kamtchatkensis Lacass., Bull. Soc. Hist. Nat.
Tsuga diversifolia on Honshu, while Betula ermanii is Toulouse 58: 637. 1929; Picea jezoensis (Siebold &
the most common broad-leaved tree. On Hokkaido Zucc.) Carrire f. kamtchatkensis (Lacass.) S. L. Tung
it occurs also in mixed coniferous-broad-leaved & Y. L. Chou, in Y. L. Chou, Lign. Fl. Heilongjiang:
forests. 46. 1986.
Picea austromandshurica Silba, J. Int. Conifer
Uses Preserv. Soc. 6 (2): 35. 1999.
Distribution Conservation
IUCN: LC Etymology
slightly convex, 1.31.9 1.11.6cm at mid-cone; sur- to arboreta where some may be mis-identified as the
face smooth, slightly striated, exposed part of abax- similar P. koyamae from Japan.
ial surface shining, glabrous; upper margin entire or
erose-denticulate, obtuse, rounded or truncate; base 2 varieties are recognized:
cuneate, dark brown. Bracts rudimentary, ligulate,
24mm, entirely included. Seeds ovoid-conical, 34
22.5mm, dark brown; seed wings ovate-oblong, Picea koraiensis Nakai var. koraiensis. Types: North
1216 68mm, pale yellowish, transparent. Korea: V. Komarov 82; T. Nakai 1880; T. Nakai s.n.
(syntypes LE, TI).
Taxonomic notes
Picea intercedens Nakai, J. Japan. Bot. 17: 4. 1941; 593
This species is very similar to Picea koyamai and Picea koraiensis Nakai var. intercedens (Nakai)
also resembles P. obovata; with the latter introgres- Y. L. Chou, Lign. Fl. Heilongjiang: 49. 1986.
sive hybridization is likely to occur. Schmidt-Vogt
(1977) recognized three varieties, described by Nakai Description
(op. cit.) as species, of these only one is sufficiently
different to be treated as a separate taxon. Seed cones short cylindrical, seed scales more or
less thick, obovate-oblong; upper margin entire,
Distribution rounded or obtuse.
successfully introduced in arboreta in the cooler or 2.6 11.7cm at mid-cone; abaxial surface smooth
colder NE of the USA and in northern Europe, but it or striated, glabrous; upper margin rounded obtuse,
has not been grown commercially. entire, or undulate, denticulate to repand-lacerate
at apex; base cuneate. Bracts rudimentary, ligulate,
2mm long, entirely included. Seeds ovoid-conical,
Picea likiangensis (Franch.) E. Pritz., Bot. Jahrb. 24mm long, dark brown; seed wings ovate-oblong,
Syst. 29: 217. 1900. 714mm long, light brown.
The species epithet refers to the Lijiang Shan in This is a highly variable species and several infraspe- 595
Yunnan, China, from where it was first described. cific taxa have been recognized. Most of these were
at one time described as distinct species and there is
Vernacular names apparently no strict consensus as to which entities to
recognize at what rank [e.g. Farjon, 1990: 287; Flora
Likiang spruce; li jiang yun shan (Chinese) of China 4: 2930 (1999)].
Description Distribution
Trees to 50m tall, d.b.h. to 22.6m; trunk mono- China: S Qinghai, S & W Sichuan, NW Yunnan, SE
podial, straight; bark rough and scaly, fissured, grey, Xizang [Tibet]; Bhutan.
with orange-brown freshly exposed bark. Branches TDWG codes: 36 CHC-SC CHC-YN CHQ CHT 40
of first order long, slender, spreading or ascend- EHM-BH
ing; branches of second order variable, not pendu-
lous; crown pyramidal or conical. Branchlets thick, Ecology
rigid or slender, firm, pale yellowish grey or orange-
brown to reddish brown; with prominent ridges Both the type (type locality Lijiang Shan, Yunnan)
and fine grooves, pubescent or glabrous; pulvini and the varieties are subalpine spruces (2700m to
11.5mm, at 6090 to shoot axis. Vegetative buds 4100m a.s.l.) of the SW Plateau of China. They are
ovoid-conical or conical, 46 34mm, (slightly) usually associated with other conifers, e.g. Abies spp.,
resinous; bud scales small, triangular, appressed, Picea brachytyla, Larix potaninii, and Tsuga spp. at
red-brown or purplish brown, persisting several the lower elevations.
years. Leaves spreading radially, forward above
shoot, parted below, (0.6)0.81.5(1.7)cm long, Uses
11.5(2)mm wide, linear, straight or curved, rigid,
quadrate-rhombic to transversely rhombic in cross- Likiang spruce is a timber tree used for construc-
section; apex acute, pungent; amphistomatic, with tion, machines, poles, furniture, and wood pulp for
fewer lines of stomata above than below; leaf colour the paper industry. The bark is used to produce tan-
dark green or glaucous green above, bluish green nin, resin is tapped or distilled from the wood, and
below. Pollen cones axillary, 22.5cm long, rose-red the needles produce aromatic oils. In Europe and
at first, yellowish at maturity. Seed cones terminal, North America this species and its varieties can be
erect at first, pendulous at maturity, often numerous, found growing in arboreta and botanic gardens, as
sessile or short, obliquely pedunculate, oval-oblong, well as in large private gardens with tree collections.
with oblique base and obtuse apex, (4)712(15)cm The correct naming to variety of these trees is often
long, (3)3.55cm wide with opened scales; colour problematic.
(immature)magenta to red, ripening to yellow-
ish brown, reddish brown, purplish brown or pale 4 varieties are recognized:
brown. Seed scales obovate or broadly obtrullate, 1.5
Conservation Distribution
Distribution
elevated it to species rank, which is here followed. Picea lutzii Little, J. Forest. (Washington) 51: 745.
More collecting in areas outside this drainage in 1953. Type not designated.
SW Sichuan and NW Yunnan, where it is reported
by the Chinese to occur, is needed to establish with Etymology
more certainty both its distribution and taxonomic
status. This nothospecies was named after H. J. Lutz of Yale
University, who discovered it in 195051.
Distribution
Vernacular names
China: SW Sichuan, NW Yunnan (Dqn Prefecture),
598 SE Xizang [Tibet]. Lutzs spruce
TDWG codes: 36 CHC-SC CHC-YN CHT
Description
Ecology
Trees to 21m tall; trunk to 50cm d.b.h. Leaves
In SE Xizang [Tibet] this species forms almost pure slightly 4angled (less so than in P. glauca), with
forests between 3000m and 3800m a.s.l., usually whitish stomatal lines on upper surface (similar to P.
well above a mixed coniferous forest belt in which sitchensis). Seed cones (3)47cm long; seed scales
Picea spinulosa is the dominant spruce. At around short and more or less rounded, thin, with erose-
3000m it occurs also mixed with Pinus armandii, denticulate upper margin, light brown.
while at its upper limit it grows with Larix sp. and
Abies spp., the firs ultimately replacing the spruces Taxonomic notes
above 36003800m a.s.l. Understorey trees include
Betula szechuanica, B. utilis, Acer caudatum, Malus A natural hybrid between Picea glauca and P. sitch-
baccata and Sorbus sp. and there may be a well devel- ensis, occurring where the parental species are
oped shrub layer with e.g. Rhododendron, Euonymus, sympatric.
Lonicera, Borinda, and Enkianthus (Rushforth, 2008).
Distribution
Conservation
USA: Alaska (Kenai Peninsula).
IUCN: LC TDWG codes: 70 ASK
Uses Ecology
This species is a timber tree used for construction, This taxon occurs in a transition between marine-
machines, poles, furniture and wood pulp for the coastal conifer forest dominated by Picea sitchensis
paper industry. The bark is used to produce tan- and (upland) interior spruce forest dominated by
nin, resin is tapped or distilled from the wood and P. glauca. Two of the commonest conifers to occur
the needles produce aromatic oils. In Europe and with it are Tsuga heterophylla and T. mertensiana, the
North America this species could be found grow- latter more in the interior of the Kenai Peninsula.
ing in arboreta and botanic gardens from earlier,
misidentified introductions. More recently, Keith Conservation
Rushforth has introduced this species in the UK
from several visits to Xizang [Tibet) between 1995 IUCN: NE
and 2000. The identification given to trees of earlier
introduction that belong to this species, which were Uses
possibly introduced by Francis Kingdon-Ward from
the Zangbo (Tsangpo) Valley, is most likely Picea This hybrid taxon is obviously being logged and
likiangensis. used as its parental species as and where it occurs.
It has also attracted some attention from forest- matal bands below. Pollen cones axillary, often very
ers in plantation experimental gardens or plots crowded, 11.5cm long, yellowish brown. Seed cones
outside North America, e.g. in the UK and in SW terminal or subterminal, often in great abundance,
Greenland, where several other northern species in clustered in top of tree, short, obliquely pedunculate
the Pinaceae are being tested for plantation forestry. or sessile, ovoid or (sub)globular; base oblique or
Hybrids between species can grow more vigorously curved, (1.5)23.5(4) cm long, 1.52(2.8) cm
than either of the parents. wide with opened scales; colour (immature) reddish
or (dark) violet, maturing to shiny red-brown or
dark purple, old cones dark red-brown, grey-brown
Picea mariana (Mill.) Britton et al., Prelim. Cat. or blackish brown, usually persisting several years
Anth. Pter. New York: 71. 1888. Abies mariana on the tree, finally deciduous. Seed scales obovate- 599
Mill., Gard. Dict., ed. 8: Abies No. 5. 1768. Type not suborbicular, rigid, brittle at last, 0.71.2 0.61cm
designated. at mid-cone; surface usually quite rough, shining,
striated or wrinkled, glabrous; upper margin erose-
Etymology denticulate, more or less undulate, usually curved
inward; base short, broad. Bracts rudimentary, ligu-
The species epithet means belonging to Marius. late, 12mm, entirely included. Seeds ovoid-cune-
ate, 2mm long, blackish brown; seed wings ovate,
Vernacular names 58mm long, orange-brown.
1.31.8 11.5cm at mid-cone; surface striated or Picea maximowiczii Regel ex Mast. var. maximo-
grooved longitudinally, dull or shiny, often resinous, wiczii. Type: Japan: Honshu, [locality not stated],
glabrous; upper margin rounded or obtuse, entire or Tschonoski (Chnosuka Sugawa) s.n. (holotype K).
slightly erose; base cuneate. Bracts rudimentary, lig-
ulate, obtuse, 23mm long, entirely included. Seeds Description
ovoid-oblong, (2.5)34.5 (1.5)2.53mm, dark
brown or grey-brown; seed wings ovate-oblong, Leaves (0.8)11.3(1.6)cm long. Seed cones (3.5)
710 45mm, yellowish brown or orange-brown. 46.5(9?)cm long; seed scales broadly obovate with
a cuneate base and a rounded, entire or slightly erose
Distribution upper margin. Seeds 34.5mm long, 2.53mm wide.
602
Japan: Honshu (Chichibu, Yatsugatake, Akaishi). Distribution
TDWG codes: 38 JAP-HN
Japan: Honshu (Chichibu, Yatsugatake, Akaishi).
Ecology TDWG codes: 38 JAP-HN
Uses
Picea maximowiczii Regel ex Mast. var. senanensis
This small bushy tree has little value for timber and Hayashi, Ill. Useful Trees (Forest Trees): sub f. 43.
is now protected from further exploitation. In Japan 1969. Type not designated.
it is commonly planted in gardens, especially in
Buddhist temple grounds, where it is valued for its Description
dense habit and slow growth; these traits also make
it a good but uncommon species for bonsai grow- This variety differs from var. maximowiczii in its
ing. Introductions to Europe and North America shorter needles (0.61.3cm) and smaller cones (2.5
have mainly been of the var. senanensis, or per- 4.5cm long), which have smaller, slightly pointed
haps involved hybrids between the two varieties. (obtuse) seed scales and smaller seeds (2.53
In European horticulture it is not a specially val- 1.52mm). Some doubt has been raised as to its tax-
ued spruce and is mainly confined to arboreta and onomic distinction (Yamazaki, 1995).
similar collections of planted trees. No cultivars are
known of this rare species. Distribution
from plantations. The wood is used for house build- of 23 lines above, 2 bands of 45 lines below; leaf
ing and other construction, foot bridges, poles, fur- colour green or dark green, with fine, greenish white
niture making, and also, especially the plantation lines of stomata. Pollen cones axillary, 11.5cm long,
timber, for wood pulp used in industrial manufac- yellow. Seed cones terminal, erect at first, pendulous
turing. This species is in cultivation for afforestation at maturity; peduncles short, oblique, or cones ses-
and as an ornamental tree in arboreta, parks, and sile; shape ovoid-oblong, tapered at base, obtuse at
large gardens, in China and (as a garden tree only) apex, (4)57cm long, 2.53cm wide with opened
in Europe and North America. scales; colour (immature) (greenish ) red or purplish
green, light brown or dull brown when ripe. Seed
scales obovate or suborbicular, opening wide, 1.21.5
604 Picea morrisonicola Hayata, J. Coll. Sci. Imp. 11.2cm at mid-cone; surface smooth or finely
Univ. Tokyo 25 (19): 220. 1908. Type: Taiwan: striated, glabrous; upper margin entire, rounded or
Nantou, Chia-i Pref., Yu-Shan, [Mt. Morrison], truncate; base broad cuneate. Bracts rudimentary,
T. Kawakami & U. Mori 2108 (lectotype TI). ligulate, purplish, 23mm long, entirely included.
Fig. 192, 193 Seeds ovoid-oblong, 33.5mm long, dark brown;
seed wings ovate-oblong, 810 45mm, light yel-
Etymology lowish or orange-brown.
606
Plate 24. Picea neoveitchii. 1. Habit of tree. 2. Branchlet with leaves. 3, 4. Seed cones. 5. Seed scale with
seeds. 6. Seed scale. 7. Leaves. 8. Seeds.
many areas illegal or unregulated timber extraction persisting several years. Leaves spreading radi-
is common. Some subpopulations are down to one ally around shoot, directed forward, parting below,
or a few trees only. 0.82.5cm long, 11.8mm wide, linear, straight
IUCN: CR [B2ab (I, ii, v); C2a (ii)] or curved, quadrangular in cross-section, acute at
apex; amphistomatic, with 24 lines of stomata on
Uses each face; leaf colour (dark) green. Pollen cones
in leaf axils, near end of shoots, 11.5cm long, yel-
The wood of this timber tree is or was used locally low. Seed cones terminal, at first erect, pendulous
for construction, poles, furniture, and wood pulp in at maturity, sessile, cylindrical, rarely ovoid-oblong,
the paper industry. From E. H. Wilsons herbarium 48cm long, 2.54cm wide with opened scales;
specimens, which include ripe cones, no trees seem colour (immature) green or red, maturing to 607
to have been grown, and its existence in cultivation brown or dark brown. Seed scales obovate-oblong
outside China remains uncertain. to flabellate, 1.52.5 12cm at mid-cone; surface
smooth, glabrous; upper margin obtuse or rounded,
entire; base cuneate. Bracts rudimentary, ligulate,
Picea obovata Ledeb., Fl. Altaica 4: 201. 1833. Picea 23mm, entirely included. Seeds ovoid-oblong,
abies (L.) H. Karst. var. obovata (Ledeb.) Lindq., 24mm, dark brown or blackish brown; seed
Acta Horti Berg. 14 (8): 307. 1948; Picea abies (L.) wings ovate-oblong or cuneate, (6)1015mm long,
H. Karst. subsp. obovata (Ledeb.) Hultn, Svensk light brown.
Bot. Tidskr. 43: 388. 1949. Type: Russia: Altai Mts.,
C. F. von Ledebour et al. s.n. (holotype LE). Taxonomic notes
Ecology Etymology
Picea obovata is a constituent of the boreal taiga of Omorika is the local name, used as the species epi-
northern Russia and Siberia, where it tends to domi- thet by Pani under Pinus.
nate on shallow soils over permafrost and occurs to
well within the Arctic Circle. In the southern parts Vernacular names
of its huge range it is forming almost pure forests or
mixed with Abies sibirica in the Altai Mountains to Serbian spruce; Morika, Omorika (Serbian)
ca. 2000m a.s.l. In water-logged areas it becomes
a stunted, narrowly columnar tree and often grows Description
608 together with Larix gmelinii in the eastern part of its
range. Betula and Populus are common associated Trees to 3040m tall, d.b.h. to 0.81m; trunk
angiosperm trees in the more or less open conifer monopodial, straight or curved at base; bark thin,
forests on slightly deeper and better drained soils. rough and scaly, with papery flakes, (reddish)
In dryer soil situations Pinus sylvestris can grow brown. Branches of first order numerous, short,
with the spruces, too. Picea obovata is extremely slender, curved downward or pendant, assurgent at
tolerant of low winter temperatures, withstanding end; branches of second order slender, pendulous,
extremes below 60 C with a totally frozen soil. hanging in dense sprays; crown narrowly coni-
Under such extreme conditions, with short but often cal or columnar, branches reaching to the ground.
hot and dry summers, it grows very slowly and trees Branchlets short, slender, flexible, orange-brown or
with a trunk diam. of 10cm can be a century or dull light brown, later grey-brown, with flat ridges
more old. and shallow, faint grooves, pubescent in 1st and 2nd
year, then glabrous; pulvini small, to 1.5mm, nearly
Conservation glabrous. Vegetative buds ovoid-conical, acute, lat-
eral buds often numerous, 58 2.54.5mm, not
IUCN: LC resinous or resinous at base; bud scales ovate, acute,
appressed, apices becoming free, red-brown, turning
Uses orange-brown, persisting several years. Leaves radi-
ally spreading at first, but soon more flat and pecti-
Siberian spruce is a major timber tree in Russia nate, with lower leaves spreading at nearly 90 from
and represents the largest resource of standing tim- shoot, (0.8)12(2.2)cm long, 1.52.2mm wide,
ber by volume in that country (and perhaps in the linear, flattened, keeled on both sides; apex obtuse
world). Much of the wood is processed to pulp, but or acutish; epistomatic, stomata in two bands of 46
other uses for which Picea wood is traditionally lines on lower surface; upper surface dark glossy
valued also apply to P. obovata, including the care- green, sometimes glaucous green, two greenish
ful construction of violins. In amenity planting it is white stomatal bands below. Pollen cones axillary,
less prominent and only used commonly in parts cylindrical, 22.5cm long, yellowish. Seed cones
of Russia and rarely in other countries of eastern terminal, often on small lateral shoots, erect at first,
Europe. pendulous at maturity; peduncles oblique or curved,
scaly; cones ovoid-oblong, oblique at base; apex
tapering, 46.5cm long, 23cm wide with opened
Picea omorika (Pani) Purk., Oesterr. Monatschr. scales; colour (immature) dark purple, maturing
Forstwesen 27: 446. 1877. Pinus omorika Pani, to purplish brown, ripe cones red-brown or grey-
Neue Conif. Alp.: 4. 1876. [Gard. Chron., ser. 2, 7: brown. Seed scales suborbicular, thin, rigid, convex,
620. 1877]. Type: Serbia: W Serbia, Rastiste, Crvena remaining imbricate for a long time, 11.5 11.3cm
Stena, J. Pani s.n. (lectotype K, designated here). at mid-cone; surface striated, sometimes transversely
undulate, often covered with dark resin, glabrous;
upper margin erose-denticulate. Bracts rudimen-
tary, ligulate, 23mm long, entirely included. Seeds
short pubescent; pulvini small, apices lighter than predominant. At lower elevations it occurs scattered
shoot. Vegetative buds ovoid-conical, acutish; termi- in broad-leaved forests, with Fagus orientalis, Acer
nal buds acute, 35mm long, not resinous; bud scales spp., Ilex colchica and Taxus baccata.
triangular, obtuse, appressed, but apices more or less
free, reddish brown, persisting several years. Leaves Conservation
spreading radially, but pressed against shoot above,
parted and nearly pectinate below, directed slightly IUCN: LC
forward, 0.60.8(1)cm long, 0.71mm wide, lin-
ear, straight, transversely rhombic in cross-section, Uses
obtuse or obliquely acutish at apex; amphistomatic,
610 12 lines of stomata on each face above, 25 lines on Oriental spruce is an important timber tree in the
each face below; leaf colour dark glossy green above, Caucasus, where it forms extensive pure stands,
whitish stomatal bands below. Pollen cones axillary, many of which are managed for forestry. It has also
12cm long, yellowish. Seed cones terminal, erect at been introduced as a forestry plantation tree in
first, pendulous at maturity, often very numerous, countries in the eastern Mediterranean. The wood
sessile, narrowly cylindrical, tapering towards apex, of this species is of good quality, comparable to that
(4.5)59(10)cm long, 23.3cm wide with opened of Norway spruce, and is put to similar uses. Among
scales; colour (immature) green, purplish green or these are construction, flooring, carpentry, furni-
purple, ripening to red-brown or dark (purplish) ture making, and parts of musical instruments. In
brown. Seed scales broadly obovate or suborbicular, horticulture, this spruce is sometimes grown as a
thin but rigid, slightly convex, 1.21.7 11.4cm at Christmas tree, but more commonly as an amenity
mid-cone; surface finely striated, shining, undulate tree for parks and large gardens in many European
or flat, sometimes resinous, glabrous; upper mar- countries and in the USA. A good number of culti-
gin rounded, entire or lacerate; base cuneate. Bracts vars is in the trade, among which are dwarf forms,
small, ligulate, serrate at rounded apex, 56mm forms with yellowish flushing leaves and those
long, entirely included. Seeds ovoid, pointed at apex, with mounding habits. The dense foliage with
34 22.5mm, red-brown or dark brown; seed very small leaves, the bright red immature pollen
wings ovate, 68 45mm, orange-brown or yel- cones, and the pendulous, purple (when still closed)
lowish brown. seed cones make this an attractive species for large
gardens.
Distribution
Caucasus, N Turkey (coastal mountains). Picea pungens Engelm., Gard. Chron., ser. 2, 11:
TDWG codes: 33 NCS-KB NCS-KC NCS-SO TCS-AR 334. 1879. Type: USA: Colorado, Clear Creek, G.
TCS-GR 34 TUR Engelmann s.n. (lectotype MO).
Ecology Etymology
Picea orientalis occurs in the mountains around the The species epithet alludes to the sharp pointed
eastern shore of the Black Sea, at elevations between (pungent) needles.
(400)7002100m a.s.l., in Turkey mainly on the
northern (seaward) slopes. It has a preference for Vernacular names
acid soils. The climate is characterized by cool to
cold winters (according to elevation) and relatively Blue spruce, Colorado spruce
warm, dry summers, the annual precipitation var-
ies between 1000mm and 2000mm; Picea occurs Description
generally on drier sites than Abies. It forms exten-
sive pure stands, especially at higher elevations and Trees to 4050m tall, d.b.h. to 11.5m; trunk mono-
at the limit of trees, or is mixed with Abies nordma- podial, straight; bark rough and scaly, deeply fis-
nniana. The undergrowth is poor, Vaccinium being sured at lower part of trunk, dark grey-brown.
Branches of first order numerous, moderately long and 3300 m a.s.l., commonly along mountain
or short, spreading horizontally, more erect or assur- streams or on moist, N-facing slopes. The soils are
gent near top; branches of second order dense, rigid, mountain lithosols and streambed gravels of vari-
spreading horizontally; crown conical, in old trees ous origin, usually poorly developed. The climate
broad columnar below the top, branches commonly is continental, with long, cold and snowy win-
to the ground. Branchlets slender, firm, yellowish ters (frost free days 60 or less) and short, but rela-
brown to orange-brown, darkest above, later turn- tively warm summers. Annual precipitation ranges
ing grey, prominently ridged and grooved, glabrous; from 600mm to 900mm. This species grows in
pulvini small, directed forward, slightly darker than small, scattered groves, especially near perennial
shoot. Vegetative buds ovoid-oblong, obtuse, 58 streams, or scattered and mixed with Pinus con-
34mm, not resinous; bud scales triangular, obtuse, torta, Pseudotsuga menziesii var. glauca, or Populus 611
loosely appressed, apices recurved, yellowish brown tremuloides. It is everywhere a rare constituent of the
or pale brown, basal scales keeled, acute, persistent, subalpine forest.
forming collars at base of shoots. Leaves spreading
radially, rigid, directed upward and forward, parted Conservation
below shoot, 1.53cm long, 11.5mm wide, linear,
slightly curved, quadrangular in cross-section, with IUCN: LC
4 prominent ribs, with acute to spinescent (pun-
gent), colourless apex; amphistomatic, 36 lines of Uses
stomata on each (grooved) face; leaf colour green,
glaucous green or bluish green. Pollen cones axillary, Blue spruce is widely planted as an ornamen-
cylindrical, 23cm long, yellow. Seed cones termi- tal; perhaps the most famous trees of this species
nal, erect at first, pendulous at maturity, numerous, are planted in front of Lenins mausoleum in the
concentrated in top of tree, sessile, ovoid-oblong or Kremlin in Moscow. Its symmetrical crown and
cylindrical, obtuse or truncate at apex, 58(10)cm glaucous blue foliage make it an ideal tree for gar-
long, 34.5cm wide with opened scales; colour dening and landscaping. In nature trees with green
(immature) green, ripening to yellowish brown or foliage exist, but these are not much wanted as the
pale brown. Seed scales obtrullate or broadly rhom- stock for horticulture; instead blue forms have been
bic, very thin, papery but tough, undulate, spread- repeatedly selected. A substantial number of culti-
ing wide, 1.82.4 11.5cm at mid-cone; surface vars, both tree forms and dwarf forms, have been
smooth, finely striated, glabrous; upper margin raised displaying various colours and hues of the
very erose; apex undulate to emarginate; base cune- needles which tend to change with their age and
ate. Bracts rudimentary, ligulate-cuspidate, 35mm become less intense. It is also popular as a Christmas
long, entirely included. Seeds ovoid, pointed at apex, tree, especially in the USA. The wood of this species
3 2mm, brown; seed wings obovate, 69 56mm, is brittle and knotty because it retains its branches
much shorter than the seed scale, yellowish on the bole and is therefore of little commercial
brown. value.
Distribution
Picea purpurea Mast., J. Linn. Soc., Bot. 37: 418.
USA: Rocky Mountains, mainly in SE Idaho, 1906. Picea likiangensis (Franch.) E. Pritz. var.
Wyoming, Utah and Colorado, with isolated popu- purpurea (Mast.) Dallim. & A. B. Jacks., Handb.
lations in Montana, Arizona and New Mexico. Conif.: 334. 1923. Type: China: Sichuan, Min River,
TDWG codes: 73 COL IDA MNT WYO 76 ARI UTA Songpan, [ad Sung Pan prope Tibetam],
77 NWM E. H. Wilson 3026 (holotype BM).
Ecology Etymology
Picea pungens is a subalpine species occurring in The species epithet refers to the purple colour of the
the Rocky Mountains at elevations between 1800m seed cones.
Purple-cone spruce; zi guo yun shan (Chinese) This species is quite unlike P. likiangensis (Franch.)
Pritzel, to which it has been linked as a variety in
Description several works describing (Chinese) conifers (e.g.
Dallimore & Jackson, 1966; Wright, 1955; Den Ouden
Trees to 4050m tall, d.b.h. to 12m; trunk mono- & Boom, 1965). It is undoubtedly closely allied to
podial, straight; bark on trunk rough and scaly, grey- P. likiangensis and its varieties and has been placed
brown, with freshly exposed parts of bark orange. with that aggregate group in the subsection Sitchenses
Branches of first order spreading horizontally, with E. Murray. The leaves of the Chinese members of this
612 lower branches curved down; branches of second subsection of spruces vary from nearly equifacial
order short, slender, dense, spreading or more pen- amphistomatic (P. likiangensis var. likiangensis) to
dant; crown pyramidal or narrowly conical, trees at nearly invers-dorsiventral epistomatic (P. purpurea).
high elevations making narrow spires. Branchlets
slender, flexible, very numerous, pale (pinkish) yel- Distribution
low-grey, finely grooved and ridged, young shoots
densely pubescent; pulvini small, directed forward China: Gansu, E Qinghai, Sichuan, NW Yunnan.
at 3050 from shoot. Vegetative buds conical, ca. TDWG codes: 36 CHC-SC CHC-YN CHN-GS CHQ
4 3mm, resinous; bud scales triangular, obtuse,
appressed, shiny dark chestnut brown, persisting Ecology
several years. Leaves above shoot closely appressed,
covering it entirely, with lower leaves more or less Picea purpurea is a subalpine species of continental
parting, all directed forward, with leaves on shaded mountains, occurring in a spruce belt at elevations
shoots more remote and spreading, 0.71.4cm long, between 2600m and 3800m a.s.l., predominantly
1.51.8mm wide, linear, straight or curved at base on N-facing slopes. The soils are either grey-brown
only, more or less dorsiventrally flattened, keeled mountain soils or lithosols, usually podzolic. The
on both sides, obtuse-mucronate at apex; usually climate is cold continental, with low to moderate
epistomatic, with two bands of densely set stomata precipitation, much of it as winter snow. It grows
below, but sometimes 12 intermittent lines of sto- in pure forests or mixed with several other species
mata above; leaf colour bright green or glossy dark of Pinaceae, e.g. Picea asperata, P. wilsonii, Larix
green above, two greenish white stomatal bands potaninii, and Abies fargesii, which prevails above
below. Pollen cones axillary, conical, 1.52.5cm long, the spruce belt towards the tree line, and with some
light red, ripening to rose or yellowish. Seed cones broad-leaved trees, usually Populus spp. and Betula
terminal, erect at first, pendulous at maturity, ses- spp. in clearings. At lower elevations Tsuga chinensis
sile, ovoid to ovoid-oblong, tapering to base, obtuse and Quercus spp. may occur.
at apex, 2.55(7)cm long, 1.73cm wide with
opened scales; colour violet, purple or bright crim- Conservation
son when immature, ripening to purplish brown
or dull (light) brown. Seed scales rhombic, papery IUCN: NT
at apex, curved, undulate, spreading wide when
ripe, 11.5 0.81.2cm at mid-cone; abaxial surface Uses
smooth or finely striated, often resinous, glabrous;
upper margin thin, elongated, undulate, incurved This species yields high quality timber used for
in some cones, erose-denticulate; base shortly nar- construction, furniture making, poles, machine
rowed. Bracts rudimentary, ligulate, 12mm long, and instument making, including musical instru-
entirely included. Seeds ovate-oblong, pointed at ments, and to a limited extent for pulp in indus-
base, 2.53mm long, (dark) purplish brown; seed trial manufacturing, e.g. paper. It was introduced to
wings (obliquely) ovate-oblong, 57 34mm, England early in the 20th century by Ernest Wilson
orange-yellowish. and Joseph Rock and is commonly found growing
in arboreta both in Europe and North America, but ward, (1)1.21.8(2.5) cm long, (1.2)1.52 mm
sometimes misidentified as P. likiangensis, or treated wide, linear, curved, quadrangular or transversely
as a variety of it (for the U.K. presumably based on rhombic in cross-section, with prominent ribs; apex
its treatment in Dallimore & Jacksons Handbook, pungent; amphistomatic, on upper surface 2 bands
1966). According to Rushforth (1987) introductions of 23 lines, on lower surface 2 bands of 46 lines of
by Wilson from W Sichuan grow to taller, more stomata; leaf colour light green or glaucous green.
columnar trees than those from Rocks collections, Pollen cones axillary, 2 35cm long, reddish, ripen-
originating from S Gansu, where the climate is drier ing to reddish yellow. Seed cones terminal, erect at
and colder in winter. first, then pendulous, sessile, oval-oblong to cylin-
dric-conical; apex obtuse, 813cm long, 2.54cm
wide with opened scales; immature cones purplish 613
Picea retroflexa Mast., J. Linn. Soc., Bot. 37: 420. red, maturing to purplish or reddish brown, ripen-
1906. Picea asperata Mast. var. retroflexa (Mast.) ing to lustrous brown. Seed scales obovate-oblong
W. C. Cheng, in Chen, Taxon. Chin. Trees: 38. or slightly obtrullate, those near base suborbicular,
1937; Picea aurantiaca Mast. var. retroflexa (Mast.) spreading wide when ripe, 1.52 1.21.5cm at mid-
C. T. Kuan & L. J. Zhou, Fl. Sichuan. 2: 71. 1983. cone; abaxial surface striated, shining, more or less
Type: China: Sichuan, Daxue Shan, near Kangding, convex, glabrous; upper margin rounded or obtuse,
[Tachien-lu], E. H. Wilson 3030 (holotype A). slightly erose-denticulate, incurved, straight or
slightly reflexed when opened base cuneate. Bracts
Etymology ligulate-lanceolate, 56mm long, entirely included.
Seeds ovoid-oblong, 34mm long, dark brown or
The species epithet (Latin retroflexus = bent back, red-brown; seed wings obovate-oblong, 1015
reflexed) refers to the pulvini on the shoots, which 57mm, pale brown or yellowish brown.
Masters described as being patenti-reflexi.
Taxonomic notes
Vernacular names
In Flora of China 4: 28 (1999) this species has been
Dapao Shan spruce; lin pi yun shan (Chinese) made a synonym of P. asperata var. asperata, while
other (Chinese) works (e.g. Ying et al., 2004; Farjon,
Description 1990, 1998, [2001]) have maintained the species rank,
or included it as a variety of P. aurantiaca (Flora of
Trees to4045m tall, d.b.h. to 11.5m; trunk mono- Sichuan). A re-examination of relevant collections
podial, straight; bark on trunk rough and scaly, and populations seems desirable; this should include
breaking into small, flaking plates, grey or brownish work on DNA sequences.
grey; inner bark yellowish. Branches of first order
short, numerous, spreading horizontally; branches Distribution
of second order short, rigid, numerous, spreading
laterally; crown narrowly conical or columnar, espe- China: W Sichuan, Qinghai (Banma Xian), S Gansu
cially in trees at high altitude. Branchlets short, firm, (Jone Xian).
thick, light brown or orange-brown, prominently TDWG codes: 36 CHC-SC CHN-GS CHQ
ridged and deeply grooved, glabrous or often ferru-
ginous pubescent; pulvini strongly developed, 12 Ecology
11.5mm, on strong shoots almost erect. Vegetative
buds broadly conical, closely surrounded by curved Picea retroflexa is a typical subalpine species, which
leaves, 510 510mm, resinous, often pubescent occurs between 3000m and 4000m a.s.l. (to 4700m
at base; bud scales triangular, keeled, appressed, E of Dawu, Schmidt-Vogt, 1977), mainly on N-facing
orange-brown or with purplish apex, persisting sev- slopes on acidic soils. The climate is continental
eral years, leaving broad collars of perular scales at alpine with low annual precipitation. At the highest
shoot bases. Leaves spreading radially, curved for- elevations it grows either pure or mixed with Abies
Etymology Distribution
The species epithet means red and describes the E Canada: maritime provinces, extreme SE Ontario,
colour of the cones (when young?). S Quebec; USA: New England States and Appalachian
Mountains.
Vernacular names TDWG codes: 72 NBR NFL-SP NSC ONT PEI QUE
75 CNT MAI MAS NWH NWJ NWY PEN VER WVA 78
Red spruce, Eastern spruce; Epinette rouge (French) NCA TEN VRG
Picea prostrata Isakov, Fl. Kirgiz. S.S.R. 10: 374. 1962. IUCN: LC
Picea sitchensis (Bong.) Carrire, Trait Gn. very thin, papery, 11.5 0.61cm at mid-cone; sur-
Conif.: 260. 1855 [sitkaensis]. Pinus sitchensis face finely striated, undulate, sometimes with dark
Bong., Mm. Acad. Imp. Sci. Saint-Petersbourg, sr. spots, glabrous; margins often recurved; apex irreg-
6, Sci. Math. 2: 164. 1832. Type not designated. ularly dentate or lacerate. Bracts small, ligulate lan-
Fig. 198 ceolate, 58mm long (sometimes nearly half as long
as seed scale), included, but often visible with wide
Etymology opened seed scales. Seeds ovoid, 23.5 1.53mm,
light or dark brown; seed wings ovate-oblong, 610
The species epithet means from Sitka a small port 45mm, light yellowish.
in Alaska, USA.
Distribution 617
Vernacular names
Pacific Coast Region of North America: from Alaska
Sitka spruce to California.
TDWG codes: 70 ASK 71 BRC 73 ORE WAS 76 CAL
Description
Ecology
Trees to 6085(90)m tall, d.b.h. to 44.5(5)m;
trunk monopodial, straight, old trees buttressed at Picea sitchensis occurs from tidewater up to steep
base; bark on trunk scaly, breaking into rough plates, mountain sides in Alaska and British Columbia,
dark grey; inner bark brown. Branches of first order generally to ca. 900m a.s.l. (highest record 1189m),
long, spreading horizontally; branches of second always in proximity to oceanic weather. The soils
order long, slender, spreading or pendant; crown are variable, usually with a thick humus layer. The
pyramidal or broad conical, in old, large trees with climate is very humid, annual precipitation ranges
abundant reiteration. Branchlets slender, flexible, from 1300mm to 3750mm, the winters are moder-
pale brown to almost white on the underside, turn- ate (in coastal Alaska snow in winter usually stays
ing yellowish or orange-brown, shallowly ridged and only above 200m). On Vancouver Island and on the
grooved, shiny, glabrous; pulvini 11.5mm, at nearly Olympic Peninsula in Washington this spruce attains
90 from shoot. Vegetative buds ovoid-conical, its greatest size. It is usually mixed with Tsuga hetero-
acute or obtuse, 45 23mm, slightly resinous at phylla (shade tolerant competitor), Pseudotsuga men-
base or not resinous; bud scales triangular, obtuse, ziesii and Thuja plicata, other associated conifers are
appressed, light brown, persisting several years Chamaecyparis lawsoniana (locally), Xanthocyparis
and becoming dark brown. Leaves spreading radi- nootkatensis, and Abies amabilis, at higher elevations
ally, stiff, on higher branches pressed forward above replaced by Tsuga mertensiana or A. lasiocarpa;
shoot, pectinate below the shoot, 1.52(2.5)cm Alnus rubra alongside rivers and Acer macrophyllum
long, ca. 1mm wide, narrowly linear, usually curved, in groves are common broad-leaved trees.
almost quadrangular in cross-section, keeled, pun-
gent at apex; stomata mainly on two (invers dorsal) Conservation
faces below, none or a few faint lines above; leaf
colour dark or bright green, with two silvery white IUCN: LC
stomatal bands. Pollen cones axillary, crowded,
23.5cm long, rose-red at first, yellowish at matu- Uses
rity. Seed cones terminal, erect at first, later pendu-
lous; peduncles short, oblique or curved; shape of Sitka spruce grows to the largest tree of its genus and
cones cylindrical-oblong, obtuse at apex, (4.5)5 is abundant in the coastal forests between roughly 43
9(10)cm long, (1.5)23(4)cm wide with opened and 62 N along the Pacific Ocean. It is a highly valu-
scales; colour (immature) yellowish green or green, able timber tree with growth rates exceeding those of
ripening to pale brown or yellowish brown. Seed other species and, in old growth stands, truly mag-
scales rhombic to obtrullate, sometimes irregular, nificent sizes. It is (still) heavily logged in clear cuts
from natural stands including old growth (in this forward at 45 from shoot, brown. Vegetative buds
part of the world this means: forest that was never ovoid conical, (5)812mm long, resinous; bud
cut before). Smaller sizes go to the paper industry, scales triangular, obtuse, keeled at base, appressed,
but big trees are prized for construction and special slightly recurved at apex, shining chestnut brown,
uses such as small aircraft, masts and spars for sail- persisting several years. Leaves spreading radially,
ing ships, oars for rowing boats, ladders, and sound- directed obliquely forward, slightly incurved, esp. on
ing boards of musical instruments. Sitka spruce has coning shoots, 2.54.5(5)cm long, ca. 1mm wide,
been widely used in plantation forestry on poor acid narrowly linear, straight or curved, soft, rhombic to
soils in cool and wet climates such as the hills and broadly rhombic in cross-section, obliquely acute or
moors of Ireland and Scotland; this timber is used acuminate at apex; amphistomatic, with 35 lines of
618 for pulp wood. In horticulture it finds less use; most stomata on each face; leaf colour dark green. Pollen
plantings in large parks as specimen trees date from cones axillary, 23cm long, yellow. Seed cones ter-
the 19th century, and only a limited number of cul- minal, erect at first, soon pendulous, obliquely short
tivars has been produced, mostly dwarf forms. It pedunculate or nearly sessile, cylindro-conical or
requires a cool and moist climate. broad fusiform (when closed) often widest near
base when opened, obtuse at apex, (8.5)1017(
18)cm long, 46cm wide with opened scales; colour
Picea smithiana (Wall.) Boiss., Fl. Orient. 5: 700. (immature) green or purplish green, maturing to
1884. Pinus smithiana Wall., Pl. Asiat. Rar. 3: 24, t. shiny bright green, ripening to lustrous brown, old
246. 1832. Type: India: Himalaya, W. S. Webb & [?] cones dull grey-brown. Seed scales obovate-fla-
Govan 6063 (lectotype C). Fig. 199, 200, 201 bellate, slightly convex, coriaceous, spreading very
wide in opened cones, 23 1.52.4cm at mid-cone;
Picea smithiana (Wall.) Boiss. var. nepalensis Franco, abaxial surface finely striated, smooth, lustrous,
Enum. Fl. Pl. Nepal 1: 26. 1978. often resinous; upper margin entire, rounded or
slightly obtuse, incurved; base cuneate. Bracts rudi-
Etymology mentary, ligulate, 45mm long, entirely included.
Seeds ovoid-oblong, pointed at apex, 57 34mm,
This species was named after James Edward Smith dark (red-) brown; seed wings ovate-oblong, 1320
(17591828), founder and first President of the 79mm, orange-brown.
Linnean Society of London.
Distribution
Vernacular names
Afghanistan: Hindu Kush; China: Xizang (Tibetan
Indian spruce, West Himalayan spruce, Morinda Himalaya); W Himalaya: Himachal Pradesh,
spruce; rai, rewari, salla, ragha, morinda (Himalaya); Karakoram, Kashmir Himalaya, Uttar Pradesh, Nepal
chang ye yun shan (Chinese) TDWG codes: 34 AFG 36 CHT 40 NEP PAK WHM-HP
WHM-JK WHM-UT
Description
Ecology
Trees to 50(60)m tall, d.b.h. to 1.52.5m; trunk
monopodial, straight; bark becoming rough, scaly, Picea smithiana is a high mountain species, occur-
greyish brown to grey, breaking into irregular plates. ring in the Himalayas in a belt between (2300)2500
Branches of first order long, slender, spreading hori- 3600(3750)m a.s.l. on alpine lithosols. The climate
zontally, often assurgent at ends; branches of sec- is moist monsoon, with abundant precipitation in
ond order long, slender, dense, strongly pendulous; two rainy seasons, but becoming gradually drier
crown conical, old trees broad columnar. Branchlets in the western parts of the range. It usually occurs
long, slender, flexible, pale yellowish brown or grey- with Abies spectabilis, Pinus wallichiana and Tsuga
ish brown, prominently ridged and grooved, gla- dumosa (eastern part of the range) and with Abies
brous; pulvini well developed, 1.5mm long, directed pindrow and Cedrus deodara in the western part.
This species occurs in the cloud belt zone of the Tigertail spruce; Torano momi, Tora momi
E Himalayas, at elevations between 2400m and (Japanese)
3700m a.s.l. (in S Xizang [Tibet] 2900m to 3600m).
It grows on alpine lithosols. The climate is moist Description
monsoon, with moderate to high annual precipita-
tion (1000mm to 2000mm), decreasing from south Trees to 3040m tall, d.b.h. to 11.3m; trunk mono-
to north. It is usually associated with Pinus walli- podial, straight; bark on trunk rough and scaly,
chiana, Larix griffithii, Abies densa, Tsuga dumosa, flaking, fissured, pale grey, with pale brown inner
620 Taxus wallichiana, and Juniperus indica; at the bark. Branches of first order long, spreading hori-
upper forest limit dense growth of Rhododendron zontally or slightly ascending; branches of second
spp. replaces the conifers. At lower elevations vari- order relatively short, dense, spreading or assurgent;
ous broad-leaved trees become more abundant: crown broad pyramidal, domed or flat topped in
Acer spp., Juglans regia, Fraxinus spp., Sorbus spp., old trees. Branchlets thick, very stout, shining yel-
Quercus pachyphylla, Prunus spp., and tree-like spe- lowish brown, later grey, prominently ridged and
cies of Rhododendron are the most common. deeply grooved, glabrous; pulvini strongly devel-
oped, 23mm long, thickened at base, spreading at
Conservation 7090 from shoot. Vegetative buds ovoid or ovoid-
oblong, obtuse or acute, 812 48mm, not or only
IUCN: LC slightly resinous; bud scales ovate, obtuse, smooth,
shining chestnut brown, appressed, persisting sev-
Uses eral years, leaving thick collars of imbricate perular
scales around shoot bases. Leaves spreading radially,
No local uses have been recorded of this species, curved, assurgent above shoot, very rigid, (1)1.5
although it is likely that its timber is being used for 2(2.5)cm long, 1.82(2.5)mm wide, linear, curved
building houses. It is quite rare in cultivation despite or twisted, quadrate-rhombic in cross-section, very
considerable garden merit; it was introduced to pungent to spinescent at apex; amphistomatic, with
England from Sikkim in 1878. It seems to have failed 46 lines of stomata on each face; leaf colour dark,
from earlier sendings of seed via the botanic garden lustrous green. Pollen cones axillary, 33.5cm long,
in Calcutta and is apparently not growing in Central reddish at first, yellow at maturity. Seed cones termi-
Europe, where the winters may be too cold. nal, erect at first, pendulous at maturity, sessile, ovoid
or ovoid-oblong, obtuse or tapering to a point (when
closed) at apex, (5)810(12)cm long, 47cm
Picea torano (Siebold ex K. Koch) Koehne, wide with opened scales; colour (immature) green
Deutsche Dendrol.: 22. 1893. Abies torano Siebold or yellowish green, ripening to cinnamon brown
ex K. Koch, Dendrol. 2 (2): 233. 1873. Type: Japan: or reddish brown. Seed scales cuneate-obovate to
[Owari pr., Abies No. 1 torano ki], P. F. von semi-orbicular (near base of cone), opening very
Siebold s.n. sub Abies polita (lectotype L). Pl. 25 wide, 22.8 1.52.5cm at mid-cone; surface finely
striated, lustrous, glabrous; upper margin entire or
denticulate, rounded, convex, sometimes lacerate;
Abies polita Siebold & Zucc., Fl. Japon. 2 (2): 20, t. base cuneate. Bracts small, ligulate-linear, 67mm
111. 1842 (nom. illeg., Art. 52.1); Picea polita (Siebold long, entirely included. Seeds ovoid, 57 34mm,
& Zucc.) Carrire, Trait Gn. Conif.: 256. 1855. brown or grey-brown; seed wings obovate-oblong,
1518 710mm, orange-brown.
Etymology
Distribution
The species epithet repeats one of the vernacular
Japanese names of this species. Japan: Honshu, Kyushu, Shikoku.
621
Plate 25. Picea torano. 1. Habit of tree. 2. Branch with foliage. 3. Green seed cone. 4. Ripe seed cone.
5. Leaf. 6. Seed.
TDWG codes: 38 JAP-HN JAP-KY JAP-SH Picea wilsonii Mast., Gard. Chron., ser. 3, 33: 133.
1903. Type: China: W Hubei, E. H. Wilson 1897
Ecology (holotype BM). Fig. 202
Picea torano occurs in (low)mountains at elevations Picea watsoniana Mast., J. Linn. Soc., Bot. 37: 419.
between 400m and 1500(1850?)m a.s.l., almost 1906; Picea wilsonii Mast. var. watsoniana (Mast.)
invariably on podzolic soils of young volcanic rocks Silba, Phytologia 68: 46. 1990.
such as lava flows and tuffs. The climate is cool, Picea wilsonii Mast. var. shanxiensis Silba, Phytologia
moist maritime, with annual precipitation exceed- 68: 46. 1990.
ing 1000mm, the winters are cold and snowy, espe-
622 cially at the higher elevations. There are some small Etymology
remnants of pure stands left, e.g. at the northern end
of Lake Yamanaka, elsewhere it is mixed with Abies This species was named after Ernest H. Wilson
homolepis, Larix kaempferi, Pinus densiflora and/or (18761930), the famous plant hunter who travelled
broad-leaved trees, e.g. Betula, Fagus, Acer, Quercus in China, Korea and Japan.
mongolica var. grosseserrata, Prunus maximowiczii,
and Zelkova serrata. Vernacular names
The more accessible and large trees have been cut in Description
the past and were replaced by afforestation with dif-
ferent species, e.g. Cryptomeria japonica and Larix Trees to 4050m tall, d.b.h. to 1.31.5m; trunk
kaempferi. monopodial, straight; bark on trunk rough and
IUCN: VU (A2ce, 3ce) scaly, greyish brown or dark grey. Branches of first
order slender, spreading horizontally; branches of
Uses second order short, slender, spreading, in some trees
becoming pendant; crown pyramidal and dense in
Tigertail spruce is uncommon and grows in inacces- young trees, columnar in old trees. Branchlets slen-
sible places, therefore it is not an important timber der, thin, flexible, pale (yellowish) grey or buff grey,
tree. In Japan it is a popular amenity tree, used in finely ridged and grooved, glabrous; pulvini very
gardens and parks. Despite its striking foliage and small, at 5070 from shoot. Vegetative buds ovoid,
cones and conical habit in cultivation, it remains an obtuse, 68mm long, not resinous; bud scales ovate,
uncommon tree in gardens and arboreta in Europe obtuse, appressed, shining, dark brown or purplish
and is even less common elsewhere outside Japan. brown, persisting several years. Leaves spreading
Despite its praise sung in earlier conifer handbooks radially, above shoot more or less imbricate, directed
(e.g. Dallimore & Jackson, 1966), a recent book on forward, parted below, (0.8)11.8(2.2)cm long,
cultivated conifers in the USA (Bitner, 2007) does 11.7mm wide, linear, straight or slightly curved,
not even mention it. It is perfectly adapted to most mostly transversely rhombic in cross-section, obtuse
climate zones in the temperate parts of the world or acute at apex; stomata on all sides, 12 lines on
and should be grown much more often; it is there each face above, 24 lines below; leaf colour glossy
where commercial conservatism and angst to try dark green. Pollen cones axillary, 23cm long, yel-
something off the well beaten track seems to dem- lowish when ripe. Seed cones terminal, erect at first,
onstrate itself most clearly. later pendulous, very numerous, sessile or obliquely
short pedunculate, ovoid-oblong or cylindrical, with
obtuse apex, (4)58cm long, 2.53.5(4)cm wide
with opened scales; colour (immature) light green
or purplish green, ripening to pale brown or dull
reddish brown. Seed scales obovate-cuneate, slightly
convex, opening wide, 1.41.7 0.91.4cm at mid- the montane boreal coniferous forests of N Sichuan
cone; surface finely striated, in old cones nearly and S Gansu it is usually growing with Picea aspe-
smooth, glabrous; upper margin entire or erose- rata and P. purpurea, in the NE of its range also with
denticulate, rounded or truncate, sometimes obtuse; P. meyeri. It is as a rule much rarer than these. Betula
base cuneate. Bracts rudimentary, ligulate, 34mm albosinensis is the most common broad-leaved tree
long, entirely included. Seeds ovoid, pointed at apex, in these spruce forests.
34.5 2.53mm, dark brown; seed wings obovate-
oblong, 1215 57mm, yellowish or light brown. Conservation
Distribution IUCN: LC
623
China: Gansu, Hebei, Hubei, Shanxi, Shaanxi, W Uses
Sichuan.
TDWG codes: 36 CHC-HU CHC-SC CHN-GS Wilson spruce is an important timber tree in China
CHN-HB CHN-SA CHN-SX yielding wood for construction, railway sleepers,
mining props, carpentry, and furniture making,
Ecology as well as for pulp used in the paper (newsprint)
industry. In horticulture, it is commonly planted as
Picea wilsonii is a high mountain species occur- an amenity tree in northern China and Russia and
ring at elevations between 1400m and 3000m a.s.l. grows well in regions with long, steady winters, but
(max. 2100m in the north of its range). The soils are suffers from late frosts in more maritime climates,
mostly non-calcareous, podzolic mountain soils. especially in western Europe. In Europe and the
The climate is continental, montane to subalpine, USA it is seldom seen outside arboreta and botanic
with low precipitation and cold, long winters. In gardens.
Pilgerodendron Florin, Svensk Bot. Tidskr. 24: 132. 1930. Type: Pilgerodendron
uviferum (D. Don) Florin [Juniperus uvifera D. Don] (Cupressaceae).
Distribution Conservation
S Argentina: Chubut, Neuqun, Rio Negro, During the period of colonization by Europeans,
Santa Cruz; S Chile: Aisn, BioBio, Los Lagos, when agriculture was introduced on a large scale in
Magellanes. many lowland and colline regions of Chile, wide-
TDWG codes: 85 AGS-CB AGS-NE AGS-RN AGS-SC spread clearance of the forests caused a considerable
CLC-BI CLS-AI CLS-LL CLS-MG decline in this and other conifer species. Its value as
a timber source, although less than that of Fitzroya,
Ecology was still a sufficient factor to increase the pressure
on this tree, causing its virtual extinction in several
This species is a codominant tree along the edges areas. Continuing pressure until recently has caused 625
of bogs and swamps of the coastal mountains, at this species to be listed in the Appendix I of CITES,
altitudes between 20m and 750m a.s.l. and grow- effectively prohibiting the export of its wood.
ing with Fitzroya cupressoides and the coniferous IUCN: VU (A1cd+2cd)
shrub Lepidothamnus fonkii. It occurs also in large
and dense stands on wet mountain slopes of islands Uses
in shallow, water-logged acidic soil over metamor-
phic rocks with Fitzroya, Podocarpus nubigenus, The timber of this species has been used for build-
Nothofagus nitida, Desfontainia spinosa, and the ing and construction of local farmsteads and other
shrub Tepualia stipularis, and with Nothofagus buildings in the past. The wood is decay resistant.
betuloides and Drimys winteri in the southernmost There is still local use and trade but, due to demi-
parts of its range. In more continental locations E nished resources, much less than in the past and
of the crest of the Andes it grows near mountain export is now prohibited. Introduced to England by
lakes. Resistance to light fires assists the regenera- William Lobb in 1849, it is still uncommon in culti-
tion of this shade intolerant species, but intense fires vation and almost restricted to arboreta and botanic
will and have killed many trees. The climate is cool gardens, which is unfortunate because it grows into
maritime with extremely high precipitation in the a handsome, conical shrub and is perfectly hardy
coastal ranges. and suitable on acidic soils in all countries with a
relatively mild winter.
Pinus L., Sp. Pl. 2: 1000. 1753. Type: Pinus sylvestris L. (Pinaceae).
Pinea Wolf, Gen. Pl.: 156. 1776. Type: Pinus pinea seed scales. Seed scales persistent, obovate to oblong,
L. Strobus Opiz, Lotos 4: 94. 1854. Type: Strobus wey- thin or thick woody, attached spirally to a slender or
mouthiana Opiz [Pinus strobus L.]. Caryopitys Small, thick rachis; exposed portion (apophysis) variously
Fl. S.E. United States: 29. 1903. Type: Caryopitys edu- thickened and/or elongated, with a terminal or dorsal
lis (Engelm.) Small [Pinus edulis Engelm.]. Apinus umbo, which may be armed with a spine or prickle.
Neck. ex Rydb., Bull. Torrey Bot. Club 32: 597. 1905. Seeds usually slightly flattened, with an adnate or
Type: Apinus cembra Neck. ex Rydb. [Pinus cembra articulate membranous wing derived from adaxial
626 L.]. Leucopitys Nieuwl., Amer. Midl. Naturalist 3: part of seed scale and several times larger than seed
69. 1913 (nom. illeg.). Type: Leucopitys strobus (L.) or reduced. Seedlings with 324 cotyledons.
Nieuwl. [Pinus strobus L.]. Ducampopinus A. Chev.,
Rev. Bot. Appliq. 24: 30. 1944. Type: Ducampopinus 113 species.
krempfii (Lecomte) A. Chev. [Pinus krempfii Lecomte]
Distribution
Pinus is the classical Latin name for pines.
North America: from Yukon to Newfoundland and
Description from SE Alaska and British Columbia south into
Mexico and Central America as far as Nicaragua; in E
Evergreen monoecious trees, less commonly shrubs; and SE USA to Florida; Bahamas, Carribean (Cuba,
resin canals in wood, bark, leaves and often cones. Hispaniola). Eurasia (N Africa): Atlas Mountains of
Trunk of trees monopodial, branching in pseudo- Morocco and Algeria; from the Iberian Peninsula
whorls, branches assurgent and repeating primary through S and Central Europe to E Europe and
branching arrangement (Rauhs model). Bark fur- Turkey, Caucasus, Syria and Lebanon; Scotland (dis-
rowed or plated, to thin and scaly or thin and smooth. junct); from Scandinavia through Russia and Siberia
Shoots dimorphic, with long shoots and dwarf shoots. to Kamchatka and Sachalin; Sino-Himalayan moun-
Primary leaves (cataphylls) scale-like, enclosing win- tain system, China, Japan, Taiwan, Indochina to N
ter buds (primordial long shoots or ovuliferous stro- Sumatera; Philippines.
bili), subtending dwarf shoot buds or pollen strobili,
early deciduous or persistent. Secondary leaves (nee- Synopsis
dles) borne in fascicles of (1)25(8) on dwarf shoots;
fascicles surrounded at base by an early deciduous The genus Pinus is the only conifer genus for which
or persistent sheath of bud scales or their remnants, several comprehensive studies investigating phy-
persisting 230 years and falling as fascicles; length logenetic relationships, based on a number of dif-
of needles 2.550cm, width 0.52.5(7)mm, acicular ferent gene sequences, have been published. There
(one species lanceolate), plano-convex or triangular is also comprehensive work available on compara-
(rarely terete or flat) in cross-section, entire or serru- tive morphology, including some cladistic analyses.
late, epistomatic or amphistomatic (one species occa- Consequently, it is possible to set up a classifica-
sionally hypostomatic). Pollen cones spirally arranged tion that is informed by (if not entirely based on) a
near proximal end of new long shoots, ovoid-oblong hypothesis of phylogenetic relationships. The most
to cylindrical; consisting of a thin axis with numer- important and consistent result of these phylogenetic
ous spirally arranged, (sub-)peltate microsporophylls, analyses is the confirmation that the genus naturally
each bearing two pollen sacs containing bisaccate divides into two subgenera. This leaves all earlier
pollen. Seed cones subterminal, borne singly or more classifications that recognized more than two pri-
commonly clustered, pedunculate, maturing in sec- mary divisions as essentially artificial; the result of a
ond year, or rarely in third year, shed early or variously choice of characters rather than a reflection of evolu-
persistent, initially erect; mature cones pendulous tionary relationships. The first comprehensive formal
or spreading, opening soon or variously serotinous, classification informed primarily by phylogeny was
(obliquely) ovoid to cylindrical, 260cm long. Bracts published by Price et al. in Richardson (1998). The
conspicuous at pollination stage, not growing with second was given in my book Pines (Farjon, 2005b),
with a review of the more recent results of molecular Section Trifolius Duhamel
phylogenetic analyses. The latter classification, with a Subsection Contortae Little et Critchfield
few minor amendments, is adopted here. Species: Pinus contorta, P. banksiana,
P. virginiana, P. clausa
Genus Pinus L. Subsection Ponderosae Loudon
Species: Pinus ponderosa, P. ari
Subgenus Strobus Lemmon zonica, P. jeffreyi, P. engelmannii,
Section Parrya Mayr P. coulteri, P. sabiniana, P. torreyana,
Subsection Nelsoniae Van der Burgh P. hartwegii, P. montezumae, P. devo-
Species: Pinus nelsonii niana, P. pseudostrobus, P. douglasi
Subsection Balfourianae Engelm. ana, P. maximinoi, P. durangensis 627
Species: Pinus balfouriana, Subsection Australes Loudon
P. aristata, P. longaeva Species: Pinus palustris, P. caribaea,
Subsection Rzedowskianae Carvajal P. elliottii, P. glabra, P. cubensis,
Species: Pinus rzedowskii, P. maxi P. occidentalis, P. pungens, P. rigida,
martinezii, P. pinceana P. serotina, P. taeda, P. leiophylla,
Subsection Cembroides Engelm. P. lumholtzii, P. herrerae, P. echinata,
Species: Pinus cembroides, P. culmini P. lawsonii, P. pringlei, P. teocote,
cola, P. edulis, P. remota, P. mono P. patula, P. greggii, P. attenuata,
phylla, P. quadrifolia P. muricata, P. radiata, P. jaliscana,
Section Quinquefolius Duhamel P. tecunumanii, P. oocarpa, P. luz-
Subsection Gerardianae Loudon mariae, P. praetermissa
Species: Pinus gerardiana, P. bunge
ana, P. squamata Keys for the genus Pinus
Subsection Krempfianae Little et
Critchfield The keys for the genus Pinus follow the taxonomic
Species: Pinus krempfii classification presented above. Keys based on taxo-
Subsection Strobi Loudon nomic classifications as opposed to those based on
Species: Pinus strobus, P. albicaulis, geographic groupings have the advantage that an
P. flexilis, P. strobiformis, P. ayaca- individual plant growing outside its native range can
huite, P. monticola, P. lambertiana, still be identified with them when the provenance of
P. cembra, P. sibirica, P. peuce, P. wal- that plant is unknown. The genus Pinus has proba-
lichiana, P. bhutanica, P. armandii, bly more species planted outside their natural ranges
P. amamiana, P. koraiensis, P. pumila, than any other genus in conifers. Excluded from
P. parviflora, P. morrisonicola, P. fen- the keys are the four hybrid species Pinus densit-
zeliana, P. wangii, P. dalatensis hunbergii, P. hakkodensis, P. neilreichiana and
Subgenus Pinus P. rhaetica.
Section Pinus
Subsection Pinaster Mayr ex Koehne Key to the subgenera of Pinus
Species: Pinus pinaster, P. heldreichii,
P. brutia, P. halepensis, P. pinea, 1a. Pulvini decurrent. Leaves with 2 vascular bun-
P. canariensis, P. roxburghii, P. latteri, dles and variously positioned resin ducts, with
P. merkusii a persistent (rarely deciduous) fascicle sheath.
Subsection Pinus (syn. Subsection Sylves Seeds with an articulate, rarely adnate, wing
tres Loudon) Subgen. Pinus
Species: Pinus densata, P. yunnanen- 1b. Pulvini not decurrent. Leaves with a single vas-
sis, P. kesiya, P. thunbergii, P. tabuli- cular bundle and external (marginal) resin
formis, P. henryi, P. hwangshanensis, ducts, with a deciduous (rarely persistent)
P. luchuensis, P. taiwanensis, P. densi fascicle sheath. Seeds with an adnate, rarely
flora, P. sylvestris, P. resinosa, P. mugo, articulate, wing or wingless when free from the
P. uncinata, P. massoniana, P. tropica seed scaleSubgen. Strobus
lis, P. nigra
Key to the subsections of Subgenus Pinus Note: There is in fact much similarity (and varia-
tion) among the species of the latter two subsec-
1a. Shoots uni-nodal; buds not resinous. Leaves in tions and many species cannot be reliably assigned
fascicles of only 2 or only 3 (rarely 2 plus some 3 to their respective subsections on the basis of their
in P. brutia). Seed cones 620(25) cm long; morphology. These subsections, while supported by
seed scales thick woody, rigid (except in P. hel- phylogenetic analyses based on DNA sequence data,
dreichii); apophyses of seed scales lustrous apparently contain much homoplasy (parallel evolu-
brown (except in P. heldreichii); umbo unarmed tion or convergence) in their morphological charac-
or at most with a minute, deciduous prickle ter states. It is therefore advised to try both keys to
(except spinaceous in P. pinaster). All Old identify a species that belongs to any one of these
628 World species Subsect. Pinaster two subsections.
1b. Not this combination of character states 2
2a. Buds (slightly) resinous or rarely not resinous. Key to the species of Pinus Subsection
Leaves in fascicles of only 2 or 2 and 3 (only 3 in Australes
P. kesiya). Seed cones deciduous or persistent
for a few years but always opening soon, 210 1a. Scales of leaf fascicles deciduous, leaving the
(12) cm long; seed scales thin woody, rigid; fascicles without a sheath before falling 2
umbo unarmed or with a minute, deciduous or 1b. Scales of leaf fascicles persistent, retaining a
sometimes persistent prickle. All but 2 Old sheath at the base of the fascicles when falling
World species Subsect. Pinus 3
2b. Not this combibation of character states 3 2a. Leaves in fascicles of 3 (exceptionally 2 or 4),
3a. Bark on lower trunk of trees with small plates (15)2030(40+)cm long, pendulous
or flakes. Shoots uni-nodal. Fascicle sheaths P. lumholtzii
shorter than 10mm; leaves in fascicles of only 2, 2b. Leaves in fascicles of (2)35(6), (4)615
28(10) cm long, rigid, curved or twisted; (17)cm long, rigid or pliant, not pendulous
resin ducts in leaves medial. Seed cones (semi-) P. leiophylla
persistent, (2)38cm long, more or less seroti- 3a. New foliage shoots multi-nodal (producing
nous or sometimes opening soon in warm sun- more than 1 whorl of branches or buds in a sin-
shine; seed scales thin woody, rigid. All New gle growing season) 4
World species Subsect. Contortae 3b. New foliage shoots uni-nodal 10
3b. Not this combination of character states 4 4a. Seed cones deciduous or semi-persistent, open-
4a. Leaves in fascicles of 3 or 5, less commonly on ing soon 5
the same tree 2, 4 or 6(8), rarely only 2 (P. pon- 4b. Seed cones persistent, serotinous 7
derosa of northern populations), straight (not 5a. Leaves in fascicles of 2, some times of 3; fascicle
rigidly curved, contorted or twisted); fascicle sheaths of new leaves 912mm long
sheaths of new leaf fascicles at least 15mm long, P. cubensis
usually 2035(40) mm long; resin ducts in 5b. Leaves in fascicles of (2)34(5); fascicle
leaves medial and sometimes 12 internal. Seed sheaths of new leaves 1530mm long 6
cone scales opening soon or sometimes more 6a. Buds resinous. Leaves rigid, (1.2)1.41.8 mm
slowly (cones not serotinous). All New World wide P. caribaea
species Subsect. Ponderosae 6b. Buds not resinous. Leaves lax, drooping, 0.7
4b. Not this combination of character states 5 0.9(1)mm wide P. patula
5. Leaves in fascicles of 23, 3 only, or 3, 4 and 5, 7a. Leaves in fascicles of only 2, thick and rigid,
not of 5 only (or more than 5) and only P. glabra 1.32 mm wide. Seed cones 57(8) cm long;
and P. muricata with fascicles of 2 only; leaf umbos of seed scales with a curved spine or
resin ducts usually or predominantly internal, sharp, persistent prickle P. muricata
sometimes medial or septal but very rarely 12 7b. Leaves in fascicles of 2 and 3, only 3, or rarely
ducts external. All New World species 4 or 5, rigid, 11.6mm wide. Seed cones (5)7
Subsect. Australes 15 cm long; umbos of seed scales unarmed or
with a minute, deciduous prickle 8
8a. Apophyses of seed scales flat or slightly raised, 15a. Seed cones broadly ovoid to subglobose when
more or less equally on all sides of the cone; closed, with a (15)2035 mm long peduncle;
umbo with a minute, deciduous prickle umbo of seed scales unarmed or with a minute,
P. greggii deciduous prickle 16
8b. Apophyses of seed scales prominently to 15b. Seed cones ovoid-conical, ovoid-attenuate, nar-
extremely raised at least on one side of the cone rowly conical or oblong when closed, with a
(in most cones); umbo unarmed 9 short (maximum 20 mm long) peduncle or
9a. Buds resinous. Pollen cones yellowish before nearly sessile; umbo of seed scales with a min-
anthesis. Seed cones ovoid-oblong to ovoid- ute, deciduous or strong and sharp prickle or
attenuate P. attenuata spine 19
9b. Buds not resinous. Pollen cones pink to reddish 16a. Leaves in fascicles of 3, rarely of 4, rigid, 1.2 629
before anthesis. Seed cones ovoid, asymmetri- 1.6mm wide P. luzmariae
cal in most cones, or nearly symmetrical if the 16b. Leaves in fascicles of 3, 4 or 5, pliant, lax and/or
apophyses are only slightly raised P. radiata drooping (sometimes more rigid), 0.51.6mm
10a. Leaves (3)412(15) cm long, in fascicles of wide 17
only 2 or 2 and 3. Buds (slightly) resinous 11 17a. Seed cones deciduous, leaving basal scales on
10b. Leaves (7)1025(45)cm long, in fascicles of 2 the peduncle not falling with the cone
and 3, sometimes 4 or 5, or only 3 (if 2 and 3 or P. praetermissa
only 3, usually longer than 12cm). Buds not res- 17b. Seed cones persistent or deciduous, if decidu-
inous or resinous (if resinous, leaves usually ous, falling intact with the peduncle attached to
longer than 12cm) 14 the cone base 18
11a. Leaves in fascicles of only 2, slender, 0.71.2mm 18a. Seed cones deciduous, opening soon; apophy-
wide. Seed cones solitary or in pairs ses of the seed scales raised P. tecunumanii
P. glabra 18b. Seed cones persistent, semi-serotinous, apoph-
11b. Leaves in fascicles of 2, 3 or sometimes 4, robust, yses of seed scales nearly flat or slightly raised
11.5(2)mm wide. Seed cones solitary or more P. oocarpa
commonly in whorls of 26 12 19a. Leaves lax and drooping, very slender, (0.5)
12a. Seed cones serotinous; seed scales with promi- 0.60.8mm wide P. jaliscana
nently raised apophyses ending in a hard, sharp 19b. Leaves rigid or pliant, 11.7mm wide 20
spine P. pungens 20a. Buds (slightly) resinous. Seed cone scales thin;
12b. Seed cones opening soon or serotinous; seed umbo of scales armed with a prickle or spine
scales with slightly raised apophyses ending in 21
a small but persistent prickle 13 20b. Buds not resinous. Seed cone scales thick;
13a. Seed cones persistent, opening soon or seroti- umbo of scales unarmed or with a minute,
nous, ovoid-conical when closed; umbos of deciduous prickle 23
seed scales with a slender, curved prickle. Seed 21a. Leaves in fascicles of 35; fascicle sheaths of
wings 34 times longer than the seeds new leaves (8)1015mm long. Seed cones (lus-
P. rigida trous) dark brown P. occidentalis
13b. Seed cones semi-persistent, opening soon, nar- 21b. Leaves in fascicles of (2)3, occasionally 4; fas-
rowly ovoid or ovoid-oblong when closed; cicle sheaths of new leaves 1525mm long. Seed
umbos of seed scales with a short, stout prickle. cones light brown 22
Seed wings 23 times longer than the seeds 22a. Leaves in fascicles of 2 and (more numerous) 3.
P. echinata Seed cones deciduous, opening soon P. taeda
14a. Leaves only in fascicles of 3. Seed cones small 22b. Leaves in fascicles of 3, occasionally 4. Seed
(2)33.5(4)cm long, deciduous cones persistent, opening soon or serotinous
P. herrerae P. serotina
14b. Leaves in fascicles of (2)3, 4 or 5. Seed cones 23a. Leaves 2045cm long, in fascicles of 3, rarely 2
usually longer than 4 cm, deciduous or or 5. Pollen cones 47(8)cm long. Seed cones
persistent 15 (15)2025cm long P. palustris
23b. Leaves (7)1025(30)cm long, in fascicles of 3b. Seed cones narrowly ovoid when closed;
2, 3, 4 or 5. Pollen cones 14cm long. Seed cones apophyses of seed scales slightly raised; umbos
(3)415(18)cm long 24 unarmed or with a small prickle P. clausa
24a. Leaves in fascicles of 2 and 3. Pollen cones pur-
plish red before anthesis. Seed cones (7)915( Key to the species of Pinus Subsection
18) cm long; apophyses of seed scales raised. Pinaster
Seeds 67 mm long, with a 2030 mm long
wing P. elliottii The species in this subsection are naturally occur-
24b. Leaves in fascicles of 35 (very rarely 2). Pollen ring in the Canary Islands, the Mediterranean, the
cones yellowish green or yellow before anthesis. western Himalaya and tropical Southeast Asia. They
630 Seed cones (3)48(10)cm long; apophyses of are likely the remnant species of a past distribution
seed scales nearly flat to slightly raised. Seeds of numerous pines that bordered the northern coasts
36mm long, with a 1218mm long wing 25 of the Thetis Ocean from the (early?) Cretaceous to
25a. Bark on the trunk breaking into small plates. the Miocene (Klaus, 1988). With one exception, they
Seed cones persistent, semi-serotinous; apoph- have seed cones with hard, rigid scales of which the
yses of seed scales lustrous brown P. pringlei apophyses are lustrous brown with unarmed umbos.
25b. Bark on the trunk deeply fissured. Seed cones The exception, Pinus heldreichii, was formerly classi-
deciduous, opening soon; apophyses of seed fied as closely related to Pinus nigra and producing
scales light brown 26 hybrids with it (see e.g. Vidakovi, 1991), but phylo-
26a. Leaves (7)1015(18)cm long, mostly in fasci- gentic analyses using DNA sequence data have dem-
cles of 3, but some of 2, 4 or 5, straight or curved. onstrated that these two species end up in different
Seed cones (3)46(7)cm long; umbos of seed clades, with P. heldreichii closely related (sister to)
scales with a minute, deciduous prickle the species here classified in subsection Pinaster and
P. teocote P. nigra positioned within the clade forming subsec-
26b. Leaves 1220(25)cm long, mostly in fascicles tion Pinus (Wang et al., 1999; Geada Lpez et al.,
of 3 and 4, sometimes of 5 (very rarely of 2), 2002). For this reason, P. heldreichii is here classified
straight. Seed cones 58(9)cm long; umbos of in subsection Pinaster.
seed scales unarmed P. lawsonii
1a. Leaves in fascicles of only 3, 2030(35) cm
Key to the species of Pinus Subsection long, pliant and drooping 2
Contortae 1b. Leaves in fascicles of only 2 or 2 and sometimes
3, 625(27) cm long, rigid or in one species
1a. Leaves curved, (0.7)1.52.5(3) mm wide. pliant but not drooping 3
Seed cones solitary or in whorls of 25. Seed 2a. Fascicle sheaths of new leaves 2530mm long.
wings 1012mm long 2 Seed cones persistent, opening slowly; apophy-
1b. Leaves straight but twisted, 11.5 mm wide. ses of scales very strongly developed, conical
Seed cones solitary or in pairs. Seed wings and often recurved P. roxburghii
1520mm long 3 2b. Fascicle sheaths of new leaves 1520mm long.
2a. Pollen cones orange-red before anthesis. Seed Seed cones deciduous, opening soon; apophy-
cones ovoid but asymmetrical at base when ses of scales prominently raised but not conical
closed, opening slowly; umbos of seed scales (except sometimes in some basal scales)
with a variable, persistent prickle P. contorta P. canariensis
2b. Pollen cones yellow before anthesis. Seed cones 3a. Fascicle sheaths of new leaves 2030mm long;
asymmetric, curved when closed, serotinous; leaves thick, 1.52 mm wide. Seed cones
umbos of seed scales unarmed P. banksiana 1020(22)cm long, persistent, opening slowly;
3a. Seed cones conical, slightly oblique when umbo on scales forming a central, rigid spine
closed; apophyses of seed scales prominently P. pinaster
raised; umbos with a strong, slender prickle to 3b. Fascicle sheaths of new leaves shorter than
5mm long P. virginiana 20 mm; leaves 0.71.5(1.8) mm wide. Seed
cones shorter than 13(15) cm, deciduous or exception). Leaves may fall off singularly from older
persistent; umbos unarmed or with a minute fascicles with deciduous sheaths; for that reason it
prickle 4 is necessary to count leaves only on recent cohorts
4a. Seed scales of cones thin and flexible; apophy- of leaves with more or less intact fascicle sheaths.
ses of scales dull brown; umbos with a minute Needle leaves of pines are grouped in fascicles of
prickle. Leaves usually curved P. heldreichii 18. There is only one species with consistently sin-
4b. Seed scales of cones thick and rigid; apophyses gle leaves (P. monophylla); common numbers are 2, 3
of scales lustrous brown; umbos unarmed. and 5. However, there are numerous species in which
Leaves straight and/or twisted 5 the numbers vary on a single tree, especially those
5a. Bark on trunk breaking into large plates. native to Mexico, but also from elsewhere. Fascicles
Fascicle sheaths of new leaves short, to ca. with 23 are common, as are 35, either with pre- 631
10mm long. Seed cones broadly ovoid to sub- dominance at the lower number, or at the higher,
globose when closed. Seeds 1520(22) mm or sometimes more equally distributed. Rarely does
long, thick, with a rudimentary wing much the number exceed 5, but in a few species in Mexico
shorter than the seed P. pinea this sometimes occurs and it can run up to 8. Several
5b. Bark on trunk breaking into small or elongated species never have more than 2 and several others
plates. Fascicle sheaths of new leaves (9)10 always have 5, but those with 3 are often more likely
20 mm long. Seed cones ovoid-conical or to have some variation, either downward or upward.
oblong-conical when closed. Seeds less than It is therefore necessary to count more than a few
10mm long, with a wing at least 2 the length fascicles in many cases, if possible from different
of the seed 6 parts of the tree or, in a population, from a few trees.
6a. Seed cones persistent, serotinous; apophyses of The number of leaves in a fascicle is not a sufficient
scales nearly flat or slightly raised. Resin ducts diagnostic character of species and needs to be sup-
in leaves external (marginal). Seedlings without plemented with other information.
a grass stage 7
6b. Seed cones deciduous, opening soon; apophy- 1a. Leaves in fascicles of only 2 2
ses of scales prominently raised. Resin ducts in 1b. Leaves in fascicles of 2 or 3, or only 3 12
leaves medial. Seedlings commonly developing 2a. Shrubs or small trees. Leaves (2.3)37(8)cm
a grass stage 8 long, rigid and curved. Seed cones semi-persis-
7a. Leaves in fascicles of only 2, (6)710(12)cm tent, obliquely ovoid when closed; apophyses of
long, 0.70.8mm wide P. halepensis scales on one side of the cone (prominently)
7b. Leaves in fascicles of 2 or sometimes 3, (5)10 raised 3
18(29)cm long, 11.5mm wide P. brutia 2b. Trees. Leaves usually longer than 7cm, straight
8a. Leaves 1520cm long, 1mm wide, pliant. Seed or when curved often pliant. Seed cones either
wings ca. 4 as long as the seeds... deciduous or long persistent (rarely semi-per-
P. merkusii sistent), more or less symmetrical when closed;
8b. Leaves 1525(27)cm long, 1.5mm wide, rigid. apophyses of scales flat or raised on all sides of
Seed wings not more than 3 as long as the the cone 4
seeds P. latteri 3a. Shrubs, rarely small trees. Apophyses of seed
scales nearly flat to prominently raised
Key to the species of Pinus Subsection Pinus P. mugo
3b. Small, monopodial trees. Apophyses of seed
Counting leaves in fascicles requires a few guidelines scales all raised, most prominently on one side
to make sure one gets the correct information from of the cone, curving backward P. uncinata
it. The adult type needle leaves of pines grow from 4a. Bark on trunk with deep, longitudinal fissures.
dwarf shoots and are held in a fascicle sheath formed Umbo of seed cone scales unarmed (without a
of bud scales. In the subgenus Pinus these sheaths spine or prickle) P. thunbergii
are mostly persistent (two exceptions) and in the 4b. Bark on trunk breaking into large and/or
subgenus Strobus they are mostly deciduous (one irregular plates. Umbo of seed cone scales with
at least a minute (but possibly deciduous) gle growing season). Leaves in fascicles of only
prickle 5 3, long and thin, lax and drooping
5a. Fascicle sheaths of new leaves 1520mm long; P. kesiya
leaves brittle (breaking easily). Pollen cones red 12b. New foliage shoots uni-nodal. Leaves in fasci-
or purple before anthesis P. resinosa cles of 2 or 3, short or long but not lax and
5b. Fascicle sheaths of new leaves 515 mm long; drooping 13
leaves rigid or pliant (not breaking easily). 13a. Buds slightly resinous. Leaves 615 cm long,
Pollen cones yellow or at most tinged red 6 rigid 14
6a. Leaves thick, robust, 12mm wide 7 13b. Buds not resinous. Leaves 730cm long, rigid
6b. Leaves thin, slender, 0.61mm wide 8 or pliant 15
632 7a. Leaves 47(12)cm long; resin ducts in leaves 14a. Bark on trunk breaking into large plates. Seed
external (marginal). Seed cones (2)37 cm cones when closed ovoid, 58.5cm long. Seeds
long, dull light brown. Seeds 35mm long with 68mm long P. tabuliformis
a 1015mm long wing P. sylvestris 14b. Bark on trunk with irregular fissures. Seed
7b. Leaves (4)816(18) cm long; resin ducts in cones when closed narrowly ovoid, 46 cm
leaves medial. Seed cones (3.5)510(12) cm long. Seeds 46mm long P. densata
long, slightly lustrous light brown. Seeds 15a. Leaves (15)2030 cm long, 1.5 mm wide,
(4)68mm long with a 1525mm long wing straight and rigid. Umbo of cone scales
P. nigra unarmed P. tropicalis
8a. Leaves ca. 1 mm wide, rigid, straight. Seed 15b. Leaves 720(30) cm long, 11.2 mm wide,
cones deciduous; apophyses of seed scales dull straight, contorted or drooping, pliant. Umbo
brown P. densiflora of cone scales mucronate, or at least with a
8b. Leaves 0.61 mm wide, pliant, straight or short prickle 16
curved. Seed cones persistent; apophyses of 16a. Bark on trunk with long, vertical fissures.
seed scales lustrous brown 9 Fascicle sheath of new leaves 1520mm long.
9a. Seed cones 2.55cm long, ovoid when closed. Seed cones deciduous P. massoniana
Wings of seeds 912mm long. Leaves 712cm 16b. Bark on trunk with irregular plates. Fascicle
long P. henryi sheath of new leaves 1015 mm long. Seed
9b. Seed cones 36(8) cm long, narrowly ovoid cones persistent P. yunnanensis
when closed. Wings of seeds 1020 mm long.
Leaves (5)1017cm long 10 Key to the species of Pinus Subsection
10a. Fascicle sheaths of new leaves 510 mm long. Ponderosae
Apophyses of seed scales nearly flat; umbo
depressed with a short, persistent prickle The species in this subsection range from western
P. hwangshanensis North America into Mexico and Central America.
10b. Fascicle sheaths of new leaves 1014mm long. The group consists of a very widespread species, P.
Apophyses of seed scales mostly slightly raised; ponderosa, some Tertiary relicts now confined to the
umbo flat or pyramidal, with a minute, decidu- two Californias, and taxa that become increasingly
ous prickle or unarmed 11 diverse genetically and morphologically in Mexico
11a. Pollen cones yellow. Umbo of seed scales nearly and Central America, where they are most likely
flat, with a minute, deciduous prickle or still evolving. The relicts are most distinct, while
unarmed. Seed wings 1520mm long the evolving species are mostly variable and can be
P. taiwanensis difficult to separate on the characters used in this
11b. Pollen cones tinged red. Umbo of seed scales key. Reference to the full descriptions is here most
slightly raised, armed with a sharp, persistent necessary to arrive at a determination with confi-
prickle. Seed wings 1015mm long dence. For counting numbers of leaves per fascicle
P. luchuensis see comments under the key to the species of Pinus
12a. New foliage shoots multi-nodal (producing Subsect. Pinus.
more than 1 whorl of branches or buds in a sin-
1a. Leaves in fascicles of not more than 3, either sures. Seed cones massive, 1535cm long. Seeds
23 or only 3 2 810mm long, with a 2535mm long wing
1b. Leaves in fascicles of 3, 4, or 5, rarely 2 or more P. devoniana
than 5 5 10b. Bark grey-brown or brown, with plates or fis-
2a. Seed cones deciduous, opening soon; scales sures less deep relative to the trunk diam. Seed
thin woody; apophyses raised but not elongated cones smaller (maximum 20 cm long). Seeds
into a strongly curved spine 3 47mm long, with a 1328mm long wing 11
2b. Seed cones persistent, opening slowly; scales 11a. Seed cones solitary or in pairs, on distinct,
thick woody; apophyses extremely raised and curved peduncles; seed scales thin and flexible.
elongated into a strongly curved spine 4 Leaves very slender, pliant, 0.61(1.1) mm
3a. Buds resinous. Pollen cones red or magenta wide P. maximinoi 633
before anthesis. Seed cones 512cm long 11b. Seed cones solitary or more often in whorls of
P. ponderosa 26, on short peduncles or appearing sessile;
3b. Buds not resinous. Pollen cones yellow or with seed scales thin or thick but rigid. Leaves rigid
a purplish tinge. Seed cones (10)1225( or pliant, 0.81.3mm wide 12
30)cm long P. jeffreyi 12a. Seed cones 710 cm long; umbo unarmed.
4a. New foliage shoots multi-nodal (producing Seeds 45mm long P. douglasiana
more than 1 whorl of branches or buds in a sin- 12b. Seed cones 720 cm long; umbo obtuse but
gle growing season). Leaves thick and rigid, strongly developed, or with a small, persistent
1.92.2 mm wide. Seed cones solitary or in prickle. Seeds 57mm long 13
whorls of 25 P. coulteri 13a. Bark on trunk breaking into small plates, not
4b. New foliage shoots uni-nodal. Leaves slender, with deep longitudinal fissures. Seed cones
pliant, 1.5mm wide. Seed cones solitary, rarely solitary or more often in whorls of 36; apophy
in pairs P. sabiniana ses flat or moderately raised P. montezumae
5a. Leaves in fascicles of 35, rarely 2 6 13b. Bark on trunk breaking into elongated plates
5b. Leaves in fascicles of 5, rarely 3, 4, or 6, very and deep, longitudinal fissures. Seed cones sol-
rarely 8 7 itary or in whorls of 23; apohyses very vari-
6a. Leaves in fascicles of (2)3(4), rarely 5, able, from nearly flat to extremely raised or
2035cm long, 1.52mm wide P. engelmannii conical, especially towards the base of the
6b. Leaves in fascicles of 35, 1020(25)cm long, cone P. pseudostrobus
0.91.8mm wide P. arizonica
7a. Leaves thick, rigid, ca. 2mm wide. Pollen cones Key to the subsections (and some species) of
before anthesis yellow, not tinged pink, pur- Pinus Subgenus Strobus
plish or brown. Seed cones persistent, opening
slowly, broadly ovoid when closed The morphological characters of species in this
P. torreyana subgenus are highly diverse but also demonstrate
7b. Leaves slender, rigid or pliant, 0.61.6 mm convergent evolution of certain traits, which means
wide. Pollen cones before anthesis usually that similar character states do appear in phyloge-
tinged pink, purplish or brown. Seed cones netically not closely related species. In the classifica-
deciduous, opening soon, ovoid-oblong or tion adopted here (see Farjon, 2005b: 218224) some
ovoid-attenuate when closed 8 weight has been given to certain autapomorphies
8a. Leaves (6)1020(24)cm long 9 (characters unique in a species or lineage that do not
8b. Leaves (15)2040(45)cm long 10 inform us about their relationships). In a few cases
9a. Leaves relatively thick, rigid, (1)1.21.5 mm the subsections therefore may diverge slightly from
wide P. hartwegii those that would only recognize (DNA based) phy-
9b. Leaves slender, rigid or pliant, 0.71.1 mm logenetic characters. This approach makes it some-
wide P. durangensis what easier to use morphological characters in this
10a. Bark on trunk reddish brown to dark brown, key to the amended subsections of the subgenus
divided by very deep, longitudinal, black fis- Strobus.
1a. Fascicle sheaths of leaves persistent, not recoil- Key to the species of Pinus Subsection
ing; leaves connate, appearing as 1 but actually Balfourianae
with 3 (rarely 4) in a fascicle. Seed cones on
relatively very long, curved peduncles 1a. Leaves with small resin droplets drying as white
Subsect. specks. Umbos on seed cone scales with a
Nelsoniae: a single species P. nelsonii strong, 410mm long prickle P. aristata
1b. Fascicle sheaths of leaves deciduous, scales first 1b. Leaves without small resin droplets. Umbos with
recoiling or not; leaves free (sometimes more or a weak, 16mm long prickle or unarmed 2
less connate from the base but always distally 2a. Bark on the trunk breaking into small, irregular
free). Peduncles of seed cones much shorter plates, creating a tessellate pattern, cinnamon
634 than the cones 2 or orange-brown. Seeds 810mm long
2a. Leaves in fascicles of only 5, these persisting on P. balfouriana
the branches for more than 10 and up to 30 2b. Bark on the trunk breaking into irregular, scaly
years, forming long foxtail tufts; fascicle ridges, shallowly to deeply fissured, reddish
sheaths not recoiling and already absent in sec- brown. Seeds 58mm long P. longaeva
ond year fascicles Subsect. Balfourianae
2b. Leaves in fascicles of 15 (very rarely 6 or 7), Key to the species of Pinus Subsection
these persisting 25 years (rarely as long as 8 Cembroides
years), with recoiling or early deciduous fasci-
cle sheaths 3 1a. Usually decumbent shrubs, multi-stemmed.
3a. Leaves flat, 1.55mm wide, in fascicles of only Leaves predominantly in fascicles of 5 (very
2. Seeds with a long wing attached Subsect. rarely of 4 or 6), persisting 23 years. Seeds
Krempfiae: a single species P. krempfii 57mm long P. culminicola
3b. Leaves not flat, 0.52(2.5)mm wide, in fasci- 1b. (Small) trees with at least a short single trunk
cles of 15 (very rarely 6 or 7); if with 2 leaves before branching. Leaves in fascicles of 15
then seeds without a wing attached 4 (rarely 6), persisting (3)48 years. Seeds
4a. Bark on trunk (and branches) remaining 1018mm long 2
smooth even in large trees, exfoliating with thin 2a. Leaves in fascicles of only 1 (rarely 2), 1.22.2
flakes creating a multi-coloured pattern some- (2.5)mm diam., terete unless with 2 in a fas-
what similar to bark of planes (Platanus) cicle; fascicle sheaths not recoiling
Subsect. Gerardianae P. monophylla
4b. Bark on trunk becoming rough and scaly at 2b. Leaves in fascicles of (1)25(rarely 6),
least in large trees, exfoliating irregularly, form- (0.6)0.71.5(1.7)mm wide, not terete unless
ing multi-layered small or large plates, or with 1 in a fascicle; fascicle sheaths recoiling or
becoming fissured 5 not 3
5a. Umbos of seed cone scales terminal (at apex of 3a. Scales of fascicle sheaths recoiling before fall-
scale). Leaves in fascicles of 5 (rarely 6 or 7) ing. Integument of seeds thick and hard 4
Subsect. Strobi 3b. Scales of fascicle sheaths deciduous without
5b. Umbos of seed scales dorsal (above apex of recoiling. Integument of seeds thin, crushed
scale). Leaves in fascicles of 15 (rarely 6) 6 easily 5
6a. Leaves 512(14) cm long. Seed cones 525 4a. Leaves 24cm long, predominantly in fascicles
(27)cm long, if shorter than 8cm only partly of 2, sometimes in fascicles of 1 or 3 P. edulis
opening with thick but flexible seed scales 4b. Leaves (2)36(8)cm long, predominantly in
Subsect. Rzedowskianae fascicles of 3, but commonly of 2 or 4 (rarely
6b. Leaves (1.5)26(8) cm long. Seed cones of 5) P. cembroides
(2)2.56(7.5)cm long, always opening widely 5a. Leaves in fascicles of 2, sometimes in fascicles
with thin, flexible seed scales of 3, 0.81.1mm wide. Seed cones (2)2.54cm
Subsect. Cembroides long P. remota
5b. Leaves in fascicles of (3)4(5), rarely in fasci- are necessary and even these can be very similar; see
cles of 2 or 6, (0.8)11.5(1.7)mm wide. Seed the note at the end of this key.
cones 46cm long P. quadrifolia
1a. Seed cones nearly sessile, persistent, serotinous.
Key to the species of Pinus Subsection Pollen cones reddish or red-purple 2
Gerardianae 1b. Seed cones pedunculate, deciduous or some-
times semi-persistent, opening soon or more
1a. Leaves in fascicles of 45; 917 cm long; buds slowly. Pollen cones yellow, greenish white or
resinous. Seed cones on 1.52cm long pedun- sometimes tinged reddish at the apex 5
cles; seeds with a ca. 15mm long wing 2a. Shrubs, spreading with layering stems. Buds
P. squamata resinous. Seed cones 35cm long P. pumila 635
1b. Leaves in fascicles of only 3; 510cm long; buds 2b. Trees. Buds not resinous. Seed cones 510
not resinous. Seed cones nearly sessile; seeds (12)cm long 3
without a wing or wing rudimentary 2 3a. Leaves persisting on the branches for up to 8
2a. Seed cones 57cm long; umbos of seed scales years, 36.5cm long P. albicaulis
with a rigid prickle or spine; seeds 810 mm 3b. Leaves persisting on the branches 24 years,
long. Leaves up to 2mm wide P. bungeana 611(13)cm long 4
2b. Seed cones 1220(23)cm long; umbos of seed 4a. Seed cones broadly ovoid, 58cm long
scales unarmed; seeds 2025mm long. Leaves P. cembra
1mm wide P. gerardiana 4b. Seed cones ovoid-conical, (5)710(12) cm
long P. sibirica
Key to the species of Pinus Subsection 5a. Seed cones opening slowly and only slightly,
Rzedowskianae late deciduous to semi-persistent; seed scales
thick and rigid; seeds without a wing or wing
1a. Leaves in fascicles of 3, rarely of 4, rigid, 0.8 rudimentary 6
1.2 mm wide; scales of fascicle sheaths early 5b. Seed cones opening soon and fully, deciduous;
deciduous, not recoiling P. pinceana seed scales thin or thick; seeds with a wing at
1b. Leaves in fascicles of (3)45, lax, 0.50.8mm least half the length of the seed, usually much
wide; scales of fascicle sheaths recoiling before longer 9
dropping off 2 6a. Leaves persisting on the branches (3)56
2a. Bark on the trunk thick, deeply fissured, dark years. Pollen cones 610mm long P. flexilis
brown. Pollen cones small, 5 3mm, purplish 6b. Leaves persisting on the branches 23 years.
before anthesis. Seed cones 1015cm long, with Pollen cones at least 15mm long 7
thin scales; seeds with a long wing attached 7a. Seed cones 57cm long, ovoid to ovoid-elliptic
P. rzedowskii when closed. Leaves (3)58(9) cm long,
2b. Bark on the trunk breaking into square plates, 0.81mm wide P. amamiana
not fissured, outer bark grey. Pollen cones 7b. Seed cones 814cm long, ovoid-oblong or con-
810mm long, yellowish before anthesis. Seed ical-cylindric when closed. Leaves (5)614cm
cones massive, (15)1725(27) cm long, with long, 11.5mm wide 8
very thick scales; seeds without a wing attached 8a. Margins of seed scales recurved. Pollen cones
P. maximartinezii yellow. Young shoots occasionally pubescent
P. koraiensis
Key to the species of Pinus Subsection Strobi 8b. Margins of seed scales not recurved. Pollen
cones greenish white with a reddish apex.
The foliage characters in this subsection of pines are Young shoots always glabrous P. armandii
very similar and, however desirable in the situation 9a. Leaves drooping or pendulous, straight but
where there are no cones, as in young trees, they can- often with a sharp bend near the base. Bark on
not be used reliably as first characters in the leads in the trunk of large trees fissured longitudinally
most cases. Seed cones and sometimes pollen cones 10
9b. Leaves lax, pliant or rigid, spreading, straight 17b. Seed cones with up to 4 cm long, straight
or (slightly) curved. Bark on the trunk of large peduncles and more or less straight at base.
trees not fissured longitudinally 11 Leaves remaining on the shoots 23 years
10a. Young shoots glandular pubescent; leaves soon 19
deciduous, remaining only 1.52 years, pendu- 18a. Pollen cones 1525 mm long. Seed cones
lous, 1524cm long P. bhutanica 611cm long. Leaves 49cm long, 0.61mm
10b. Young shoots glabrous; leaves remaining on wide
the shoots 23 years, drooping or sometimes P. morrisonicola
pendulous, (6)1018(20)cm long 18b. Pollen cones 1015mm long. Seed cones (7)8
P. wallichiana 15(20) cm long. Leaves 710(12) cm long,
636 11a. Seed cones in clusters of 48(10), nearly ses- 0.50.7mm wide P. peuce
sile, persistent, ovoid or sub-globose when 19a. Seed cones (10)1540(50)cm long, 715cm
closed, 47cm long P. parviflora wide when open. Leaves (8)1015(18) cm
11b. Seed cones solitary or in whorls of 24, pedun- long P. ayacahuite
culate, deciduous, ovoid to long cylindrical, 19b. Seed cones (5)620(25)cm long, 49cm wide
variable but usually longer than 7cm 12 when open. Leaves (3)512 (14)cm long 20
12a. Seed cones with thick, rigid scales and these 20a* Buds slightly resinous [American species]
with thick, woody apophyses, large (but vari- P. strobus
able in one species) to 5560cm long 13 20b* Buds not resinous [Vietnamese species]
12b. Seed cones with thin, flexible or more rigid P. dalatensis
scales and these with thin or only partly thick-
ened apophyses, (3)540(50)cm long 14 * Both leaf lengths and seed cone lengths are extremely
13a. Seed cones when mature at least 30 cm long, variable in these two species, ranging from
with 512 cm long peduncles. Bark on large (3)512(14)cm in the leaves and (5)623(25)
trunks breaking into large plates cm in the cones. These species, with disjunct ranges
P. lambertiana in eastern North America, southern Mexico and
13b. Seed cones when mature 1230(60)cm long, parts of Guatemala (P. strobus), and Viet Nam
on 1.52.5 cm long peduncles. Bark on large (P. dalatensis), are remarkably similar and probably
trunks breaking into small plates closely related. The Mexican-Guatemalan popula-
P. strobiformis tions are by some accepted as a third species, Pinus
14a. Seed scales near base (but well above the chiapensis, but are here maintained as a variety of P.
peduncle) upturned or reflexed 15 strobus, which during the Ice Ages occupied refugia
14b. Seed scales near base (but well above the far to the south of its current range and extended
peduncle) incurved or more or less straight probably into Mexico.
and flat 17
15a. Seed cones 1025(30) cm long; seeds with a
2025mm long wing P. monticola Pinus albicaulis Engelm., Trans. Acad. Sci. St. Louis
15b. Seed cones (3)415(17)cm long; seeds with 2: 209. 1863. Type: USA: Colorado, J. S. Newberry
a rudimentary or short wing to 16 mm long s.n. (US No. 61140) (holotype US). Fig. 205
16
16a. Leaves 2.56 cm long; resin ducts in leaves Etymology
medial P. wangii
16b. Leaves 418 cm long; resin ducts in leaves The Latin epithet albicaulis means with a white
external, sometimes a few medial P. fenzeliana stem, referring to the light grey bark.
17a. Seed cones with short, curved or obliquely
inserted peduncles, or more or less curved at Vernacular names
base. Leaves remaining on the shoots 34 years
18 Whitebark pine, Scrub pine
Description Distribution
Trees to 1015(21)m tall, d.b.h. to 11.5m. Trunk W North America: Rocky Mountains, Cascade
monopodial or with 2several stems, straight or con- Range, Sierra Nevada.
torted, branching very low and much reduced in size TDWG codes: 71 ABT BRC 73 COL IDA MNT ORE
at tree line. Bark thin, rough and scaly, breaking into WAS WYO 76 CAL NEV
small, scaly plates, yellowish brown, weathering pale
grey, on young trees and branches smooth and grey- Ecology
ish white. Branches of first order contorted, usually
short, ascending or spreading, persistent, of higher Pinus albicaulis is a high altitude pine growing in
orders flexible, assurgent. Shoots often puberulent, subalpine conifer forests up to the timberline. It 637
with short, non-decurrent pulvini, pale red-brown, occurs at altitudes up to 1350m a.s.l. in the north
ageing to pale grey-brown. Cataphylls 57mm long, and 3650m a.s.l. in the south of its range. It is a very
subulate, scarious, brown, with entire margins, soon slow growing and long-lived tree, with often con-
deciduous. Vegetative buds ovoid and acute; terminal torted stems and branches on exposed mountain
bud 810mm long; lateral buds smaller, with imbri- ridges. Whitebark pine occurs in a cold, moist, and
cate, appressed, subulate, red-brown scales. Fascicle snowy climatic zone exposed to high winds, where
sheaths 812mm long, with 67 loosely imbricate, few trees can compete. Most commonly it is associ-
orange-brown scales, deciduous and absent in sec- ated with Abies lasiocarpa, Tsuga mertensiana, and
ond years fascicles. Leaves in fascicles of 5, in dense in regions with less abundant precipitation, Pinus
tufts towards ends of branchlets, persisting up to 8 contorta. Rain only occurs in the short summer
years, mostly connivent, curved forward against period (June-September), during the remainder of
shoot, 36.5cm long, 11.5mm wide, with serrulate the year sleet and snow prevail, and the snow pack
margins distally, acute, yellow-green on abaxial face, can cover small trees entirely except on windswept
white stomatal lines on both adaxial faces. Stomata: ridges. The strong, flexible branches of P. albicau-
leaves epistomatic or weakly amphistomatic, with lis can withstand winds of hurricane force. Pinus
01 lines of stomata in grooves on abaxial face and albicaulis co-evolved with a mountain bird, Clarks
23 (intermittent) lines on each adaxial face. Pollen Nutcracker (Nucifraga columbiana, Corvidae; see
cones crowded at the proximal end of a new shoot, also under P. cembra) in a mutualism concerning
spreading, subtended by light brown bracts, ovoid- seed dispersal. The bird removes the seeds from the
oblong, 1015mm long, red, maturing to brown. closed cones without damaging them and puts them
Microsporophylls peltate, the distal part cordate to away for later use in caches. Those that are forgot-
rounded, smooth, ca. 1mm wide. Seed cones subter- ten may germinate, often two to five trees thus grow
minal, solitary, in pairs or in whorls of 3, on very short close together. No germination occurs without help
peduncles, persistent. Immature cones erect, ovoid, from the bird and the bird even feeds its young with
dark grey or purplish, maturing in two seasons. the pine seeds. Red squirrels play a similar but lesser
Mature cones broadly ovoid to subglobose, 58 role in seed planting.
47cm. Seed scales ca. 50, remaining closed, con-
cavo-convex, with seed cavities near base on adaxial Conservation
side, dark grey or purple, maturing to light brown.
Apophysis thick, triangular to rhombic, transversely The most severe threat to this species at present is
keeled, straight or recurved but not reflexed at mid- infestation by the introduced rust Cronartium ribi-
cone, light brown, resinous. Umbo terminal, trans- cola, which has led to reductions of 8090% in some
verse-triangular, acute, 56mm wide, grey-brown, northern areas of its range, where there is a wetter
very resinous. Seeds obovoid, 711 47mm, chest- climate. Additional threats are predation by the bee-
nut brown. Seed wings absent when the seed is free tle Dendrococtonus ponderosae (attacking especially
from the scale. weakened pines infested by the rust) and fire sup-
pression in National Parks leading to a successional
change in vegetation. Attempts are being made to
select rust-resistant genetic variants and to adjust cataphylls ovate-linear, reddish brown. Leaves in
the fire regimes in National Parks to a more natural remote fascicles of 5, held by a soon deciduous sheath
frequency. This species has a very wide distribution of flimsy scales, persisting 23 years, slender and
and not all populations are similarly effected. spreading, (3)58(9)cm long, straight or slightly
IUCN: EN (A4a, c, e) curved, triangular in cross-section, 0.81mm wide;
stomata in lines on the two adaxial faces only. Pollen
Uses cones in long clusters, spirally arranged at base of new
shoots, cylindrical or ovoid-ellipsoid, 1.52.5(3)cm
Whitebark pines main value is providing habitat long, slender or stout, greenish white with a slightly
for wildlife. The wood is used locally to build log reddish apex. Seed cones solitary or in pairs, initially
638 cabins and for firewood. The edible seeds are diffi- erect and green, becoming more or less pendulous
cult to harvest due to the serotinous and resinous on stout, curved, 12cm long peduncles, opening
cones; they were used by Native Americans only only slightly, ovoid to ovoid-elliptic, 57cm long,
as an emergency food source. Its aesthetic value is 34cm wide when opened, often resinous, turn-
best appreciated in situ in the high mountains; the ing dark purplish brown. Seed scales woody, rigid,
horticultural merits of this pine are low because of 1.52cm long, 23cm wide, curved slightly inward;
extremely slow growth and vulnerability to patho- basal scales scarcely or not recurved, on the adaxial
gens such as white pine blister rust (Cronartium side with two marked depressions holding the seeds.
ribicola, Basidiomycota) and late frost in parks and Apophysis rhombic or more or less rounded distally,
gardens. It is therefore scarcely known in cultivation. thick woody, with a straight or slightly recurved edge
terminating in an inconspicuous, obtuse umbo, light
yellowish brown or darker red-brown. Seeds ellip-
Pinus amamiana Koidz., Bot. Mag. (Tokyo) 38: soid, 1012mm long, 56mm wide, ca. 4mm thick,
113. 1924. Pinus armandii Franch. var. amamiana grey to nearly black, wingless or with a small ridge
(Koidz.) Hatus., Mem. Fac. Agric. Kugos Univ. 1: on abaxial margin, rarely with a rudimentary wing.
37. 1974. Type not designated.
Taxonomic notes
Etymology
The relationship of this species with the very
This species was named after the Amami people, similar species P. armandii, P. fenzeliana and
who inhabit the Ryukyu Islands. P. morrisonicola and the infraspecific taxa in this
group of East Asian pines belonging to subsection
Vernacular names Strobus is in need of further investigation using
DNA sequence data. Pinus amamiana may be most
Amami pine; amami-goyo, yaku-tane-goyo distinct in its cones and seeds, which seem to be
(Japanese) more morphologically adapted to seed dispersal by
birds, with small cones and relatively large, virtually
Description wingless seeds. However, as we have learned from
DNA analysis of other pines with such cones (e.g.
Trees to 20(30)m tall; trunk to 2m d.b.h. Bark P. albicaulis and P. cembra) strong selective pressure
smooth in young trees, greyish brown, becoming has determined the evolution of these adaptations
fissured and breaking into large and flaking plates and they are not necessarily good indicators of phy-
on lower trunk, grey or blackish grey. Branches in logenetic relationships.
pseudo-whorls in young trees, forming a conical
crown; in older trees crown widening and becoming Distribution
rounded and open. Foliage branches smooth, young
shoots puberulent, soon glabrous, grey-green, turn- Japan, Kyushu (Yakushima, Tanegashima).
ing grey-brown. Buds cylindrical, slightly resinous; TDWG codes: 38 JAP-KY
terminal bud 1015mm long; lateral buds smaller;
Pinus amamiana occurs in exposed, open stands in Rocky Mountain bristlecone pine, Colorado bristle-
often sparsely vegetated localities on rocky slopes cone pine, Hickory pine
at between 50m and 900m a.s.l. on two smaller
islands in the south of Japan. Description
640
Plate 26. Pinus arizonica var. arizonica. 1. Habit of tree. 2. Leaves. 3. Seed cones, one green, one ripe.
1220 48mm, semi-transparent, greyish brown cabinetwork, boxes, and woodware. There are many
with black tinge. local or regional sawmills where the logs are sawn
and relatively little is exported. The resource has
Distribution become much reduced by overexploitation and it
is difficult to implement more sustainable forestry
SW USA: Arizona, New Mexico; in Mexico mainly methods in the short run that would reverse the
in the Sierra Madre Occidental, southwards to S downward trend. This pine is not known to be in cul-
Durango, scattered in Coahuila, NE Zacatecas and tivation outside a few trial plantations, but perhaps
Nuevo Len. an occasional tree from southern Arizona identified
TDWG codes: 76 ARI 77 NWM TEX 79 MXE-CO as P. ponderosa planted in parks or gardens belongs
642 MXE-CU MXE-DU MXE-NL MXE-ZA MXN-SI to this species.
MXN-SO
3 varieties are recognized:
Ecology
Pinus arizonica is forming pure stands or is more Pinus arizonica Engelm. var. arizonica. Pinus
commonly mixed with Quercus spp., other pines, e.g. ponderosa Douglas ex C. Lawson var. arizonica
Pinus engelmannii, P. strobiformis, and occasionally (Engelm.) Shaw, [Pines Mexico] Publ. Arnold
Juniperus flaccida at lower or J. deppeana at higher Arbor. 1: 24. 1909; Pinus ponderosa Douglas ex
altitudes. It grows on various substrates, but the best C. Lawson subsp. arizonica (Engelm.) E. Murray,
stands are in valleys and on mesas with deep soil, Kalmia 12: 23. 1982. Type: USA: Arizona, Pima
in moderately dry to mesic forest with light winter Co., Santa Rita Mts., J. T. Rothrock 652 (holotype
frost occurring, at altitudes from (1300)20002700 MO). Pl. 26
(3000)m a.s.l. Annual precipitation is low to mod-
erate, from 700mm to 900mm, mostly falling dur- Description
ing the winter months.
Trees, usually tall, height to 3035m. Leaves in fas-
Conservation cicles of 35 (predominantly 34), rigid to slightly
lax, straight or slightly curved and twisted, (8)10
Pinus arizonica is an important constituent of the 20(23)cm long, 0.91.4(1.6)mm wide. Stomata
pine and pine-oak forests of northern Mexico, espe- on all faces of leaves, in (3)48 lines on the convex
cially in the Sierra Madre Occidental. As such it abaxial face and in (3)4(5) lines on each adax-
is heavily exploited as a timber tree in much of its ial face (abaxial number of lines correlated with
range, especially in the more accessible regions, and width of leaf). Seed cones ovoid to broadly ovoid,
stands with large trees in these localities are now often slightly curved, (4.5)57 3.56cm when
rare (Perry, 1991). While the species as a whole is open. Seeds obliquely ovoid, slightly flattened, 46
widespread and still abundant, two of its variet- 33.5mm. Seed wings obliquely ovate, 1215
ies with limited distributions are now assessed as 46mm.
Vulnerable.
Distribution
Uses
SW USA (Arizona, New Mexico); mainly in the
In Mexico, Pinus arizonica is a major timber tree, Sierra Madre Occidental of Mexico southwards to S
often compared (or equated) with P. ponderosa in Durango, scattered in Coahuila, NE Zacatecas and
the USA. The uses of its wood are similar. Timber Nuevo Len.
of large sizes and high grade is sawn for light con- TDWG codes: 76 ARI 77 NWM 79 MXE-CO MXE-CU
struction like window frames, doors, stairs, flooring, MXE-DU MXE-NL MXE-ZA MXN-SI MXN-SO
sidings, panelling, and veneers and for furniture,
figure 177. Nothotsuga longi
bracteata in Nan Ling Mts. Hunan,
China
644
figure 186. Phyllocladus
hypophyllus glaucous foliage on
Mt. Kinabalu 3100m
645
figure 210.
Pinus balfouri-
ana in the
Sierra Nevada,
California,
USA
647
figure 212.
Pinus bungeana
seed cone
figure 216.
Pinus cembroides
var. cembroides
bark
648
650
in lines on the two adaxial faces only. Pollen cones tree it is of limited value and exploited only for
in long clusters, spirally arranged at base of new local use.
shoots, cylindrical or ovoid-ellipsoid, 1.52.5(3)cm
long, slender or stout, greenish white with a slightly 3 varieties are recognized:
reddish apex. Seed cones solitary or in pairs, initially
erect and green, becoming more or less pendulous Pinus armandii Franch. var. armandii. Type not
on stout, curved, 23cm long peduncles, falling late designated. Fig. 207
after seed dispersal, conical-cylindric, 814cm long,
58cm wide when opened, often resinous, turning Description
variously brown. Seed scales woody, rigid, 34
652 2.53cm, curved slightly inward,basal scales scarcely New shoots green or grey-green. Pollen cones stout,
or not recurved, on the adaxial side with two marked 1.52cm long, ovoid-ellipsoid. Apophyses in mature
depressions holding the seeds. Apophysis rhombic cones thick woody, rhombic, not recurved or only
or triangular, often thick woody, with a straight or the umbo slightly recurved, yellowish brown or light
slightly recurved edge terminating in an incon- brown. Seeds dark brown to nearly black.
spicuous, obtuse umbo, light yellowish brown or
darker red-brown. Seeds obovoid, 1015mm long, Distribution
610mm wide, light or dark brown to nearly black,
wingless or with a small ridge on abaxial margin, China: Chongqing, S Gansu, Guizhou, Hainan,
rarely with a rudimentary wing. Henan, Hubei, Shaanxi, Sichuan, Yunnan, extreme
SE Xizang [Tibet]; N Myanmar [Burma].
Distribution TDWG codes: 36 CHC-CQ CHC-GZ CHC-HU
CHC-SC CHC-YN CHH CHN-GS CHN-SA CHN-SX
China, Anhui, Chongqing, S Gansu, Guizhou, CHS-HE CHT 41 MYA
Guangxi, Hainan, Henan, Hubei, S Shaanxi, Sichuan,
Yunnan, extreme SE Xizang [Tibet]; N Myanmar Conservation
[Burma]; Taiwan.
TDWG codes: 36 CHC-CQ CHC-GZ CHC-HU CHC- IUCN: LC
SC CHC-YN CHH CHN-GS CHN-SA CHN-SX CHS-
AH CHS-HE CHT 38 TAI 41 MYA
Pinus armandii Franch. var. dabeshanensis
Ecology (W. C. Cheng & Y. W. Law) Silba, Phytologia 68: 47.
1990. Pinus dabeshanensis W. C. Cheng &
Pinus armandii occurs in mountains at altitudes from Y. W. Law, Acta Phytotax. Sin. 13 (4): 85. 1975;
900m to 3500m a.s.l.; usually in association with Pinus fenzeliana Hand.-Mazz. var. dabeshanensis
other conifers and seldom in pure stands. Common (W. C. Cheng & Y. W. Law) L. K. Fu & Nan Li,
conifer genera in these mixed forests are Abies, Picea, Novon 7 (3): 262. 1997. Type: China: Anhui, Yuexi,
Pseudotsuga, and in SW China also Larix. More Laibangmen, C. Q. Zhang 1 (holotype PE).
often than these conifers, the pines tend to occupy
rocky areas with thin soils where other trees, among Description
them angiosperms, are less competitive.
Leaves 514cm long. Apophysis of seed scales thick
Uses woody. Seeds light brown, with a rudimentary
wing.
This pine is a notable ornamental tree in China
and it was introduced to France by Armand David Taxonomic notes
in 1895. It remains an uncommon tree in gar-
dens and parks outside China, but has spread to In Flora of China 4: 24 (1999) this taxon is treated
arboreta in many parts of the world. As a timber as a variety of Pinus fenzeliana Hand.-Mazz. Farjon
Conservation
Pinus attenuata Lemmon, Mining Sci. Press 64: 45.
This variety is known from disjunct localities in a 1892. Type: USA: California, Berkeley, California,
limited area where three provinces meet and where J. G. Lemmon s.n. (lectotype UC). Fig. 208
deforestation is known to occur and has occurred
for a long time. In Flora of China it is called an Etymology
endangered plant but it is not clear whether this is
based on IUCN criteria. The species epithet refers to the tapering (attenuate)
IUCN: VU [A2acd; B1ab (iii, v)] shape of the closed seed cone.
Vernacular names
Pinus armandii Franch. var. mastersiana (Hayata)
Hayata, J. Coll. Sci. Imp. Univ. Tokyo 25 (19): 217, Knobcone pine, Narrowcone pine
f. 8. 1908. Pinus mastersiana Hayata, Gard. Chron.,
ser. 3, 43: 194. 1908; Pinus armandii Franch. subsp. Description
mastersiana (Hayata) Businsk, Acta Pruhoniciana
68: 13. 1999. Type: Taiwan: Nantou, Chia-i Pref., Trees to 1520(25)m tall, d.b.h. to 4050cm.
Yu-Shan, [Mt. Morrison], G. Nakahara s.n., Trunk monopodial, straight or sometimes curved.
3 Oct 1905 (holotype TI). Bark thin, rough, scaly, breaking in small, rectangu-
lar plates, greyish brown to grey. Branches spread-
Description ing to assurging, forming an irregular or rounded,
open crown. Shoots often multi-nodal, with short
New shoots grey-brown. Pollen cones when fully decurrent, persistent pulvini. Cataphylls subulate,
grown slender, cylindrical, 23cm long. Apophyses scarious, tapering into a caudate apex; margins
usually slightly recurved towards apex, brown or erose-ciliate, dark brown. Vegetative buds ovoid-
red-brown when mature. Seed colour not recorded. acute; terminal bud 1530 1015mm; lateral buds
smaller, resinous, brown; the scales appressed, subu-
Distribution late, scarious, with erose, hyaline margins. Fascicle
sheaths initially 1018mm long, reduced to 36mm.
Taiwan (Mt. Alishan, Mt. Yushan). Leaves in fascicles of 3, rarely 2, persisting 23 years,
TDWG codes: 38 TAI rigid, straight, (8)1012(14)cm long, 1.01.5mm
Mexico: from Central Mexico southwards to Pinus ayacahuite Ehrenb. ex Schltdl. var. ayaca-
Chiapas; Guatemala; Honduras; El Salvador. huite. Type: Mexico: Hidalgo, Omitlan de Juarez,
TDWG codes: 79 MXC-DF MXC-ME MXC-MO Montes del Lucero, near Hacienda de Guerrero,
MXC-PU MXC-TL MXE-GU MXE-HI MXE-QU C. A. Ehrenberg 647 (lectotype MO).
MXG-VC MXS MXT-CI 80 ELS GUA HON
Description
Ecology
Seed cones pendulous, cylindrical, curved, taper-
Pinus ayacahuite forms emergent trees in mixed ing towards apex, (10)1540cm long, 715cm wide
montane conifer forest on mesic sites, or grows in when opened. Seed scales ca. 100120, thin, flex-
groups or small groves; reaching its greatest extent ible; apophysis thin, trullate, usually recurved, rarely
in Chiapas and W Guatemala. The best stands are reflexed at apex. Seeds 810 68mm. Seed wings
on loamy, well drained soils. Its altitudinal range is 2035 812mm, tapering towards apex.
(1500)19003200(3600)m a.s.l.
Pinus ayacahuite Ehrenb. ex Schltdl. var. veitchii This species was named after (but not by) John
(Roezl) Shaw, [Pines Mexico] Publ. Arnold Arbor. Hutton Balfour (18081884), who for bibliographic
1: 10. 1909. Pinus veitchii Roezl, Cat. Grain. Conif. reasons is to be credited with the authorship.
Mexic.: 32. 1857. Pinus strobiformis Engelm.
subsp. veitchii (Roezl) Frankis, Int. Dendrol. Soc. Vernacular names
Yearb. 2008. 2009. Type: Mexico: Mxico, Valle de
Mxico, Volcn Popocatepetl, [ridge between Mts. Foxtail pine
Popocatepetl and Iztaccihuatl], C. G. Pringle s.n.
(neotype A). Fig. 209 Description
to brown. Microsporophylls peltate, the distal part population) and of the southern Sierra Nevada (the
cordate to rounded, smooth, ca. 1mm wide. Seed southern populations). In the north it is found at
cones subterminal, solitary or frequently in pairs, altitudes of between 1600m and 2400m a.s.l., in the
nearly sessile, falling after seed dispersal. Immature south between 2900m and 3700m. Stands of this
cones erect, ovoid, dark purple, maturing in two pine are very open and occur on dry, rocky, exposed
seasons. Mature cones ovoid-cylindric becoming high slopes and ridges, usually devoid of other signif-
more ovoid when open, 610 46cm. Apophysis icant vegetation. Stands may be pure or mixed with
on seed scales thick, triangular to rhombic, trans- P. albicaulis, sometimes Juniperus occidentalis grows
versely keeled, recurved, red-brown. Umbo dorsal, with it, too. Regeneration and growth are extremely
transverse-triangular at base, unarmed or terminat- slow and stands commonly look as if entirely com-
ing in a weak prickle 1mm long. Seeds ellipsoid or posed of veteran trees of great age. Regeneration 657
narrowly obovoid, 810 46mm, light brown with is probably episodal and may be linked with cli-
darker red-brown specks. Seed wings 1012mm long. matic cycles. Unlike its even longer lived cousin
P.longaeva, growing only 3540 km to the east of the
Taxonomic notes Sierra Nevada, little is known about the exact ages
of some of the oldest trees, but they are likely to be
The two disjunct populations have been treated as more than 2000 years old.
distinct subspecies or varieties, with the southern
population named P. balfouriana subsp. austrina. Conservation
The differences are very small and include chemi-
cal characters (terpene compounds), which are This species may be at risk in the long term from
of doubtful significance in taxonomy. The popu- climate change if this is continuing to accelerate,
lations being disjunct, it is likely that some differ- possibly bringing in competitors and/or pathogens
ences can be found, e.g. in leaf colour, and that these it may not be able to cope with. At present, both dis-
may be partly genetic in origin. Mostly the differ- junct populations are well protected within National
ences involve character gradients and would appear Parks and National Forest Wilderness Areas and
to be not truly distinct if wide sampling was done unaffected by long-term effects of fire suppression
before analysing the data. Pinus balfouriana appears in forests as the trees usually occur in remote sub-
to be closely related to P. longaeva, a species which alpine locations where such measures have not been
occurs to the east, also in isolated populations on undertaken. It would be profitable to study the (aut-)
high mountains. There are at least some consistent ecology of this species in more detail in order to be
differences in the morphology between these two able to estimate risks under various climate change
species, but it could as well be defended that there scenarios.
is only a single species with two or three subspecies IUCN: NT
or varieties.
Uses
Distribution
Foxtail pine is not a timber tree due to extremely
USA: mainly California, in two disjunct populations slow growth and general inaccessibility of the stands
in northern (Klamath Mts.) and in southern (Sierra of relatively small trees. The dense and hard wood
Nevada) California. One locality (metapopulation) is obviously of value for special uses like wood craft
of the northern population is just across the state and some dead and down wood may be used for
border in Oregon (Lanner, 2007). this purpose. Almost all stands of this pine are now
TDWG codes: 73 ORE 76 CAL within protected areas and felling as well as dead
wood collecting are there strictly prohibited. This
Ecology species was introduced to Britain in the early 1850s
by the Scottish Oregon Association through the
Pinus balfouriana occurs in the subalpine to alpine services of John Jeffrey, who collected seeds of west-
zones of the Klamath Mountains (the northern ern American trees for the gentlemen who instituted
the association for this purpose. It is still in cultiva- depressed or sunken, unarmed or with a minute,
tion for gardens, but rare and virtually restricted to reflexed prickle. Seeds obovoid, flattened, 45mm
collections of trees and shrubs. Only two cultivars long, brown to blackish, with a 1012mm long wing.
have been named, both in the USA
Distribution
Pinus banksiana Lamb., Descr. Pinus 1: 7, t. 3. N North America, from Nova Scotia and
1803. Type: Illustration in Lambert, Descr. Pinus 1: Pennsylvania to Northwest Territories and Alberta.
t. 3. 1803. (lectotype). TDWG codes: 70 NWT 71 ABT MAN SAS 72 NBR
NSC ONT QUE 74 MIN WIS 75 CNT INI MAI MAS
658 Etymology MIC NWH NWJ NWY PEN VER
yields per ha even under natural conditions. Its otherwise straight, 1mm wide, triangular in cross-
wood is also used in carpentry and joinery, for con- section; margins minutely serrulate; leaf colour
tainers, pallets and crates as well as particle-board. bright green abaxially, glaucous white adaxially; sto-
It was introduced to Europe in 1785, but has limited mata in fine lines on the two adaxial faces. Pollen
value in horticulture and is present only as specimen cones in small clusters at base of new shoots, spirally
trees in most arboreta in countries or regions with arranged, with 812 basal perular scales remain-
a cool to cold climate. A few mutants, some from ing and partly covering the short cylindrical, yel-
witches brooms, are grown as cultivars. Extensive low cones. Seed cones solitary or in whorls of 26,
planting in forestry has usually been associated with initially erect on stout, (1)46cm long peduncles,
afforestation of poor sandy soils in northern regions; becoming pendulous when growing, cylindrical,
there it remained a small tree of irregular shape in usually slightly curved, 1220cm long, 34cm wide 659
most instances. when closed and 57cm wide when mature and
opened, usually resinous, soon falling with peduncle
after seeds have been released. Seed scales cuneate-
Pinus bhutanica Grierson et al., Notes Roy. Bot. oblong, widest just below the apophysis, thin woody,
Gard. Edinburgh 38: 299. 1980. Pinus wallichi- only slightly flexible, with two seed cavities near
ana A. B. Jacks. subsp. bhutanica (Grierson et al.) adaxial base, dull red-brown with lighter marks of
Businsk, Acta Pruhoniciana 68: 10. 1999. Type: seed wings. Apophysis rhombic, slightly raised,
Bhutan: Mongar District, Tashiyangsi, 4 km N of obtusely keeled, light brown with darker terminal
Tschilingor, A. J. C. Grierson & D. G. Long 2282 umbo. Seeds obovoid, 68mm long, 45mm wide,
(holotype E). slightly flattened, brown; wing persistent, 1522mm
long, 710mm wide, grey-brown.
Etymology
Taxonomic notes
The species epithet refers to Bhutan, from where it
was first described. This species was formerly not recognized as distinct
from Pinus wallichiana and herbarium specimens
Vernacular names belonging to P. bhutanica may remain unrecognized
as the distinction is more in the habit (strongly pen-
Bhutan pine, Bhutan white pine dulous leaves) and young shoots (not always col-
lected). Some botanists regard it as a subspecies or
Description variety of Pinus wallichiana. The identification of a
seedling in China is therefore dubious, as seedling
Trees to 25m tall or more; trunk to 80cm d.b.h., characters were not originally described from this
bole straight and columnar. Bark smooth on young species and no comparative study growing both spe-
trees and on branches of large trees, becoming fis- cies has been published.
sured and breaking into small scales that flake off in
small chips, dark greyish brown. Branches whorled, Distribution
spreading wide and sinuous, those of higher orders
drooping but with upturned ends, forming a some- Bhutan; NE India: Arunachal Pradesh (Kameng dis-
what open crown. Foliage branches slender or stout, trict); SW China: NW Yunnan, SE Xizang [Tibet].
new shoots glandular pubescent, whitish pruinose, TDWG codes: 36 CHC-YN CHT 40 EHM-AP
becoming pale green and then grey. Buds ovoid- EHM-BH
conical; terminal bud 1015mm long; lateral buds
smaller and more ovoid, slightly resinous; cataphylls Ecology
grey with an orange tinge or light reddish brown.
Leaves in fascicles of 5, held in deciduous, basal fas- The species occurs in a (warm) temperate zone,
cicle sheaths of orange-brown flimsy scales, persist- from ca. 1700m to 3050m in Bhutan, at 2100m in
ing 1.52 years, very slender and flexible, 1524cm SE Xizang [Tibet], and at 18002200m a.s.l. in NW
long, pendulous, often with a bend near the base but Yunnan, with reports down to 1000m in Arunachal
Pradesh. Associated species can include Pinus wal- times forked. Bark thin, orange-brown, becoming
lichiana, Pinus roxburghii (dry inner valleys) and thick only on lower trunk of large trees, then deeply
various broadleaved trees. It has been reported from fissured longitudinally, scaly, breaking into elongated
a variety of forest types, including secondary forest plates, pale brown to red-brown. Branches long,
where logging has occurred. spreading and ascending, forming a broad pyra-
midal or rounded, open crown. Foliage branches
Conservation slender, 35mm thick (to 10mm on cone-bearing
shoots), becoming rough with pulvini from fallen
The known range of this species (extent of occur- leaf fascicles, glabrous, new shoots glaucous green,
rence or EOO) does not meet the threshold for a yellowish brown and finally grey. Buds ovoid-con-
660 threatened category. It is also thought to occur more ical, acute, 1015mm long, not resinous; cataphylls
widely eastwards, but botanical collecting has been with recurved apices, reddish brown fringed with
virtually absent in this region since this species was white hairs. Leaves in fascicles of 2 (sometimes 3),
first described in 1980. It is reported to be harvested held by a 1016mm long, persistent sheath, retained
as part of a mixed timber resource, but there is no on branchlet 23 years, straight and rigid, spread-
evidence of decline and new localities have been ing, rarely lax and pendulous, (5)1018(29)cm
found. New collections have recently been made in long, 11.5mm wide; margins minutely serrulate;
China (Yunnan, Gaoligong Mts.) and an assessment leaf colour bright or dark green; stomata in fine lines
of the species in this area is required. on all surfaces. Pollen cones spirally arranged, short
IUCN: LC cylindrical, 12cm long, yellow. Seed cones solitary
or in whorls of 23(4), short pedunculate to nearly
Uses sessile, persistent, spreading forward or at right angle
to shoot when full grown, narrowly or broadly ovoid-
The uses as a timber tree are unknown. In horticul- conical or ovoid to rarely subglobose when closed,
ture, Bhutan pine has been introduced in the UK (4)611(13)cm long, variously serotinous, 35cm
in 1979 and it now grows well in S England, as well wide when closed, 58cm wide when opened. Seed
as in Ireland (Grimshaw & Bayton, 2009: 590591). scales thick woody, rigid, straight, oblong; apopyses
However, it is still restricted to botanic gardens and nearly flat or slightly raised, (sharply) transversely
arboreta despite its appealing habit and gracefully keeled and with thin rays radiating from the cen-
pendulous foliage. tre, more or less rhombic or often with a rounded
upper margin, to 20mm wide at mid-cone, lustrous
red-brown weathering grey; umbo flat or depressed,
Pinus brutia Ten., Fl. Napol. 1, Prodr.: lxxii. 47mm wide, broadly rhombic in outline, tan or
181115. grey-brown, unarmed. Seeds obovoid, slightly flat-
tened, 67(8)mm long, grey-brown, sometimes
Etymology dark mottled; wing 1420mm long, 811mm wide,
oblique, grey-brown with darker streaks.
The species epithet is presumably derived from
Brutium, a Roman district, now Calabria in south- Distribution
ern Italy.
E Mediterranean Region; around the Black Sea;
Vernacular names Caucasus; Turkey; NW Iran; N Iraq.
TDWG codes: 13 GRC KRI TUE 14 KRY 33 NCS-KR
Calabrian pine, Brutia pine; Kzlam (Turkey) TCS-AZ 34 CYP EAI IRN IRQ LBS-LB LBS-SY TUR
Description Ecology
Trees to 30(35?)m tall; trunk to 1.5(2.1)m d.b.h., Pinus brutia can form extensive, relatively open pine
usually a straight bole, or slightly sinuous, some- forests, either pure or mixed with Cupressus semper-
Pinus brutia has been planted extensively in coun- Pinus brutia Ten. var. eldarica (Medw.) Silba,
tries around the eastern Mediterranean and Black Phytologia 58: 367. 1985. Pinus eldarica Medw.,
Sea as it is the easiest pine to grow (with P. halepen- Trudy Tiflissk. Bot. Sada 6 (2): 21. 1903. Type:
sis) in the Mediterranean climate. It was originally Illustration in Trudy Tiflissk. Bot. Sada 6 (2): 21.
described from Calabria in Italy, which is probably a 1903 (lectotype).
planted source. Frequent use of P. halepensis sources
from the western Mediterranean threaten to destroy Description
the genetic distinctions between the two species,
possibly also in natural stands of P. brutia. The latter Seed cones ovoid-globose; apophyses slightly raised,
species has a better stem shape and growth from a whitish grey.
forestry point of view and should therefore be pro-
tected. Its timber is used for fencing posts, telephone Taxonomic notes
posts, building timbers, railway sleepers, carpentry,
boxes and crates, hardboard and pulp. The resin of Caucasian and Russian botanists usually refer to
both pines has been used from ancient times to fla- this taxon as a distinct species, Pinus eldarica; how-
vour white wines known as retsina and is still tapped ever, its distinctive characters are minor and may
especially in Turkey, now mainly for the production not warrant taxonomic recognition at all. A criti-
of turpentine. In horticulture its use is less common, cal taxonomic revision involving the species pair
mainly as an occasional amenity tree in villages and P. brutia P. halepensis is overdue and should
towns around the Mediterranean Sea; this species was include this taxon. The use of DNA sequence analy-
also tried as a forestry plantation tree in SE Australia. sis in such a study is paramount, as morphology is
not very informative in these species.
4 varieties are recognized:
Distribution
Pinus brutia Ten. var. brutia. Pinus halepensis Mill. Afghanistan?; Azerbaijan (near border with
var. brutia (Ten.) A. Henry, Trees Great Brit. Ireland Georgia); Georgia; NW Iran; N Iraq.
5: 1100. 1910; Pinus halepensis Mill. subsp. brutia TDWG codes: 33 TCS 34 IRN IRQ
(Ten.) Holmboe, [Stud. Veg. Cyprus] Bergens Mus.
Skr., ser. 2, 1 (2): 29. 1914. Type not designated. Ecology
Pinus brutia Ten. var. densifolia F. Yaltirik & M. Semi-dry pine forests, usually mixed with sclero-
Boydak, Karaca Arbor. Mag. 5 (4): 175. 2000. phyllous angiosperms in an open canopy.
Conservation Conservation
The range of this variety of P. brutia remains imper- IUCN: VU [B2ab (ii, iii, v)]
fectly known and an assessment of its conservation
status is also made difficult for lack of information
about trends at (sub)population level. Pinus bungeana Zucc. ex Endl., Syn. Conif.: 166.
IUCN: DD 1847. Type not designated. Fig. 211, 212
Etymology
Pinus brutia Ten. var. pendulifolia Frankis, in
662 Taskin (ed.), Papers Int. Symp. Pinus brutia Ten.: This species was named after the Russian botanist
14. 1993. Type: Turkey, Mugla, near Kayadibi, Alexander Andrejewitsch von Bunge (18031890).
M. P. Frankis 63 (holotype E).
Vernacular names
Description
Lace-bark pine; bai pi song (Chinese)
Leaves 1829cm long, pendulous. Seed cones ovoid-
conical; apophyses red-brown. Description
Distribution Uses
Macaronesia, Canary Islands (La Palma, Tenerife, Pinus canariensis was heavily exploited for its tim-
Gran Canaria, El Hierro). ber in the 19th and early parts of the 20th century;
TDWG codes: 21 CNY the wood was exported. Reforestation with this
species has been sucessful, but today the timber is
Ecology considered less important than the water retain-
ing capacity of the belt of pine forest especially on
In the Canary Islands this species occupies fairly Tenerife and Gran Canaria. The ever increasing
extensive forest areas above an altitude of 1200m tourism and development of built areas put a great
to ca. 2200m a.s.l. It grows well on exposed moun- demand on limited resources of drinking quality
tain slopes of volcanic origin amidst or on old lava water, which even with ample precipitation on the
flows and on scoria. Winter snow and frost limit its western islands would be quickly lost downhill and
altitudinal range upwards (Pico del Teide is 3718m); into the sea through the extensive areas of scoria
at lower elevations a thermophile woodland and/or and cavernous lava flows, if it were not for the water
xerophytic scrub vegetation, interspersed with rare retaining capacity of the pine forest belt. This spe-
evergreen lauraceous woodland on N-facing slopes cies has been introduced to Italy and South Africa
and in moist gullies, dominates. Pinus canariensis is as a forestry plantation tree. Its wood is suitable for
dominant within this pine belt where it exists, but building timbers because of its good strength, and
spring-flowering leguminose and other herbs and is widely used in light construction, carpentry, wall
shrubs may be common. Some common shrubs are panelling, turnery, and marquetry. Mass uses as for
Adenocarpus foliolosus, Chamaecytisus proliferus, pit props in mining and chipboard are now discon-
and Cistus symphytifolius. The pine woods are char- tinued, as forests are required to regrow to former
acteristically very open and sunny and often there is extent and capacity after a period of overexploita-
nothing else on the ground than rocks, pine needles tion. In horticulture it is uncommon, but present
and great numbers of the large seed cones rolled into in gardens and parks in the Mediterranean region
hollows. and in some arboreta elsewhere in countries with a
similar climate. Growing this and other pines from
warm climates under glass in cooler countries will
not succeed; pines need the movement of wind to
665
Plate 27. Pinus canariensis. 1. Habit of tree. 2. Branchlet with leaves. 3. Leaves. 4. Seed cone.
build stems that can support a crown. Specimens I basal scales, rhombic to pentagonal in outline, trans-
have seen in glasshouses beyond sapling size had to versely keeled, up to 15mm wide, ochraceous, light
be hung from the ceiling by a steel wire. or dark brown, lustrous. Umbo dorsal, flat or slightly
raised or pyramidal and curved upward, with a min-
ute, persistent prickle. Seeds obovoid, slightly flat-
Pinus caribaea Morelet, Rev. Hort. Cte dOr 1: tened, 57 2.53.5mm, light grey-brown with dark
107. 1851. spots, or dark brown to blackish. Seed wings articu-
late or adnate, obliquely ovate or oblong, 1020
Etymology 58mm, thin, semi-transparent, yellowish grey or
light brown with dark stripes.
666 The species epithet refers to its occurrence in the
Caribbean; the species was originally described from Distribution
Isla de la Juventud [Isla de Pinos] in Cuba.
West Indies: Bahamas, Turks-Caicos Islands, W
Vernacular names Cuba (including Isla de la Juventud [Isla de Pinos]);
Mexico (S Quintana Roo); N Guatemala; Belize;
Caribbean pine; pino de la costa, pino colorado, Honduras (including Islas de la Baha); Nicaragua.
pino macho (Spanish) TDWG codes: 79 MXT-QR 80 BLZ GUA HON NIC 81
BAH CUB TCI
Description
Ecology
Trees to 2035(45)m tall, d.b.h. to 50100cm;
trunk monopodial, erect. Bark rough, scaly, breaking See under varieties.
into irregularly square plates, grey-brown. Branches
spreading or ascending, of higher orders similar, or Uses
drooping. Crown broad conical, open or irregular.
Shoots multi-nodal, very rough, resinous; with large, Pinus caribaea, and especially the Mesoamerican
short decurrent and persistent pulvini. Cataphylls variety hondurensis, is an important timber tree
1520mm long, subulate, strongly recurved, scari- forming easily accessible, extensive stands in level
ous, with hyaline-ciliate margins, dark brown. to moderately hilly terrain. As such it is heavily
Vegetative buds ovoid-oblong to cylindrical; termi- exploited and although not in danger of extinction
nal bud 2025mm long, but lateral buds smaller, in any of its major areas, many local and especially
ovoid-acute, (slightly) resinous. Fascicle sheaths outlying populations which are likely to be geneti-
1520mm long, retaining their length or reduced to cally distinct are now severely degraded (Dvorak
ca. 10mm at maturity. Leaves in fascicles of (2)3( & Donahue, 1992). One of these is Ejido Caobas,
4), very rarely 5, persisting 3 years, straight, (slightly) Quintana Roo, Mexico, which is the northernmost
twisted, rigid, (12)1526(28)cm long, (1.2)1.4 occurrence of P. caribaea var. hondurensis. Pinus
1.8mm wide; margins serrulate; apex acute-pungent; caribaea has been widely planted as a timber tree
leaf colour light or dark green (occasionally glaucous in tropical and subtropical lowland areas around
green), more or less lustrous. Stomata conspicuous the world. The heavily resinous wood is used for
on all faces of leaves, in 811(14) lines on the con- building, crates, pallets, boat decks, poles and posts,
vex abaxial face and 46 lines on each adaxial face. plywood, and particleboard, as well as for pulp
Pollen cones cylindrical, often curved when mature, manufacture. The latter use comes primarily from
23cm 56mm, pink or yellow, turning yellowish plantations outside its natural range. In Honduras,
or reddish brown. Seed cones subterminal, mostly in trees have been widely tapped for resin, but this
pairs or whorls of 35(8), on 22.5cm long pedun- activity has become less common recently. This
cles, ovoid-conical or ovoid when opened, more or species is scarcely known in horticulture and can
less symmetrical but base obliquely flattened, (4)5 only be grown successfully in the subtropics and
12(13) (3)46(7)cm when open. Seed scales tropics.
oblong, straight or recurved, dark (purplish) brown
or blackish brown. Apophysis raised or nearly flat on 3 varieties are recognized:
Pinus caribaea Morelet var. caribaea. Type: Cuba: wide, light or dark green. Seed cones (4)510(12)
Isla de la Juventud [Isla de Pinos], Santa Barbara, (3)46(7)cm when open; apophysis rhombic to
H. A. Lckhoff 11608 (neotype PRF). pentagonal in outline, ochraceous or light brown,
lustrous. Cotyledons (4)67(9), 1225mm long.
Description Seedlings with an elongated stem; primary leaves
green, more or less ascending, soon replaced by sec-
Fascicle sheaths 1520mm long, retaining their ondary leaves.
length at maturity. Leaves in fascicles of 3(4), very
rarely 2 or 5, (13)1526cm long, (1.2)1.41.8mm Distribution
wide, light or dark green. Seed cones (4)510(12)
(3)46(7)cm when open; apophysis rhombic to Bahamas: on Grand Bahama, Great Abaco, New 667
pentagonal in outline, ochraceous or light brown, Providence and North and South Andros Islands
lustrous. Cotyledons (4)67(9), 1225mm long. in the north; on Great Inagua Island in the south.
Seedlings with an elongated stem; primary leaves Also on the Turks and Caicos Islands in three dis-
green, more or less ascending, soon replaced by sec- tinct subpopulations at Pine Cay, North Caicos and
ondary leaves. Middle Caicos.
TDWG codes: 81 BAH TCI
Distribution
Ecology
West Indies: W Cuba, Pinar del Ro; Isla de la
Juventud [Isla de Pinos]. On flat to rolling, eroded limestone rock with pock-
TDWG codes: 81 CUB ets of sandy alkaline soil, or on sandy spits and old
beaches, forming pure, open fire-climax stands, usu-
Ecology ally with scattered or dense undergrowth of shrubs
(often Sabal palmetto); also invading open scru-
Forming pure, open, dry fire-climax forest or open bland forming secondary forest. Altitudinal range
woodland with undergrowth of grasses or scattered 110m a.s.l. Annual precipitation decreases from
shrubs on sandy or gravelly, well-drained, acidic ca. 1500mm on Grand Bahama to ca. 1000mm on
soils. Altitudinal range from 1700m a.s.l., with South Andros in the northern Bahamas (750mm on
most extensive stands below 400m. The growing is TCI); mean annual temperature is ca. 25 C and frost
season continuous in a warm tropical climate with does not occur. Hurricanes are a recurring destruc-
long dry spells. Annual precipitation varies mainly tive factor.
with altitude, between ca. 10001800mm, with a
winter dry season. Frost does not occur. Conservation
Pinus caribaea Morelet var. hondurensis (Sncl.) Pinus cembra L., Sp. Pl. 2: 1000. 1753. Type:
W. H. Barrett & Golfari, Caribbean Forest. 23: Illustration: Larix sempervirens, foliis quinis,
65. 1962. Pinus hondurensis Sncl., Conif.: 126. nucleis edulibus in Breyn in Acad. Caes.-Leop.
1868. Type: Belize: El Cayo, San Pastor Pine Ridge, Carol. Nat. Cur. Ephem. 7: 8, t. 1. (1719). (lecto-
Chiquibul F.R., Angels Drive, W. H. Barrett 77582 type). Fig. 213, 214
(neotype BAB).
Etymology
Description
The species epithet is from the Italian vernacular
Fascicle sheaths 1520mm long, retaining their name cembro for this pine.
668 length at maturity. Leaves in fascicles of 3 (rarely 2, 4,
very rarely 5), (12)1628cm long, (1.2)1.41.8mm Vernacular names
wide, light green, occasionally glaucous green. Seed
cones (4)512(13) 3.57cm when open; apophy- Arolla pine, Swiss stone pine; Arve, Zirbel (German);
sis rhombic to pentagonal in outline with upper auvier, pin arle (French); cembro (Italian); limba
margin irregularly undulate, chestnut-brown, lus- (Polish); borovice limby (Slovak)
trous. Cotyledons 58, 2035mm long. Seedlings
with an elongated stem; primary leaves glaucous, Description
more or less spreading, development of secondary
leaves delayed. Trees to 25m tall; trunk to 1.5m d.b.h., often
crooked and stunted at high altitude. Bark smooth
Distribution and orange-brown on young trees and on branches,
becoming scaly and deeply fissured on trunks of
Mexico (S Quintana Roo); Belize; N & E Guatemala; large trees, turning grey with red-brown fissures.
Honduras (including the Islas de la Baha); Branches spreading and assurging, forming a
Nicaragua. broadly conical crown, often flat-topped in exposed
TDWG codes: 79 MXT-QR 80 BLZ GUA HON NIC trees. Foliage branches slender to stout, initially with
a dense, brown pubescence, then glabrous, smooth,
Ecology light brown. Buds small, ovoid-conical, without
resin; cataphylls reddish brown. Leaves in dense
Mainly distributed on the lowland coastal plains tufts towards end of shoots, persiting 24 years, in
within the Atlantic climatic influence, from the edges fascicles of 5 held by deciduous basal sheaths of thin,
of mangrove swamps to lower upland bunch-grass/ orange-brown scales that fall away in the second
pine savannas. It occurs on well-drained, sandy or year, spreading wide or forward, straight or slightly
gravelly, acidic soils, forming pure (or sometimes curved, more or less rigid to flexible but not lax,
mixed with P. oocarpa and/or P. tecunumanii) fire- (6)79cm long, triangular in cross-section, not
climax forest, or pine-oak forest, with undergrowth twisted, 11.5mm wide; margins minutely serrulate;
of grasses, Pteridium aquilinum, or Sabal palmetto leaf colour green; stomata in white lines predomi-
nearer the coast, possibly replacing broad-leaved nantly on the two adaxial faces, a few stomata may
rainforest under human influence in many areas. occur on abaxial side. Pollen cones clustered, spirally
The altitudinal range is 1700(1000?)m a.s.l. arranged, short cylindrical, initially reddish turning
Annual precipitation varies greatly, from the high- red-brown. Seed cones intitially erect, becoming pat-
est measurement at Laguna del Pinar in Nicaragua ent, single or in 23 in whorls on very short pedun-
(ca. 4000mm) to the lowest at Los Limones in the cles, remaining closed or opening only slightly at
interior of Honduras (ca. 660mm). Mean annual maturity, from green or glaucous green to purplish
temperature is from 2127 C and no frosts occur. when growing, resinous, broadly ovoid, 58cm long,
46cm wide, dark brown when ripe. Seed scales
Conservation imbricate, widely cuneate proximally, with 2 deep
seed cavities adaxially, soft woody. Apophysis broadly
IUCN: LC rhombic or widely triangular to semi-orbicular,
thickened; margins free or slightly reflexed; umbo it grows with European larch and the latter is much
terminal, triangular, obtuse, pale brown. Seeds more valued for timber, forestry practises tend to
oblong-obovoid, 1016 68mm, without a wing favour the latter, which in succesional terms is the
or wing rudimentary. pioneer species. The wood has been used for the
building of traditional houses and is valued for spe-
Distribution cial uses such as joinery, panelling, cabinet making,
tools, and wood turning. The seeds, though edible,
Europe: Alps, Carpathians. are difficult to harvest due to the soft, resinous and
TDWG codes: 11 AUT-AU CZE-CZ CZE-SL GER POL closed cone scales and are consequently mostly left
SWI 12 FRA-FR 13 ITA-IT ROM YUG-BH 14 UKR-MO to the birds. As an ornamental tree it is quite valu-
UKR-UK able, but it is sensible to late frosts as are other 669
conifers from very cold regions where spring means
Ecology spring and is not interrupted by a brief return to win-
ter. Despite this, a number of cultivars are known,
In the inner valleys of the Alps Pinus cembra, both with distinct habit and with divergent needle
together with Larix decidua, forms open conifer for- colours; they are particularly well grown in northern
est and woodland up to the tree line at between 2200 and eastern Europe.
and 2600m altitude. Arolla pine may descend down
to 1200m, where it is usually a rare component of
conifer forest dominated by Picea abies. Centuries Pinus cembroides Zucc., Abh. Math.-Phys. Cl.
of intensive grazing mainly with cattle have brought Knigl. Bayer. Akad. Wiss. 1: 392. 1832.
the tree line in the Alps down and turned much of
the ancient forest into pasture woodland. This veg- Etymology
etation is in places dominated by Ericaceae such as
Vaccinium myrtillus and Rhododendron ferrugineum, The species epithet means similar but not equal to
grasses and herbs. However, more recently alpine cembra, i.e. Pinus cembra L.
grazing has been substantially reduced and the for-
est is making a come-back on many slopes. Arolla Vernacular names
pine is dependent on the European Nutcracker
(Nucifraga caryocatactes, Corvidae) for its effective Pinyon pine, Mexican nut pine; pin (Spanish)
seed dispersal and the birds carry the seeds and
therewith the pines up slope. Pinus cembra is most Description
probably a Siberian element in the European flora
and is resistant to 40 frost; unlike accompany- Shrubs or trees 315(25)m tall, d.b.h. to 1080
ing the larches, P. cembra has evergreen foliage and (120)cm. Trunk monopodial, very short (shrubs)
it reduces the water content in the needles during to medium size, branching low. Bark thick, rough
winter to a minimum. Pinus cembra is slow growing and scaly, breaking into small, irregular, shaggy
and can live to great age (>1000 years in the Swiss plates, grey or greyish brown. Branches spreading or
Aletschwald) having rot-resistant wood. trailing on the ground, or ascending. Crown an open
or dense shrub or broad, open, irregular. Shoots
Conservation orange-brown or slightly glaucous at first, soon grey.
Cataphylls small, 24mm long, subulate or trian-
IUCN: LC gular, acute-acuminate, with erose margins, light
brown, caducous. Vegetative buds ovoid-oblong to
Uses oval-cylindrical; terminal bud 58(10) x 35mm;
lateral buds smaller, not or slightly resinous, light
Arolla pine is not a significant timber tree due to brown or ochraceous. Fascicle sheaths short,
its slow growth and commonly curved or contorted 46mm long, loosely imbricate, the scales (pale)
shape, although the tree can grow quite straight brown, soon recoiling, then pale straw-coloured to
and to considerable size in protected localities. As grey, forming a small rosette at base of fascicle but
deciduous before the leaves fall. Leaves in fascicles and valleys and more mesic coniferous montane
of (2)3(4, rarely 5), persisting (3)45(7) years, forests. The altitudinal range is extensive: (800
curved or less often straight, lax or sometimes rigid, )15002600(2800)m a.s.l., with highest occur-
(2)36(8)cm long, (0.6)0.71(1.2)mm wide; rences in the SE of its range. It grows on a variety of
margins entire; apex acute-acuminate or pungent; substrates, ranging from alluvial bajadas to volcanic
leaf colour variable, dull green to glaucous green, rock, usually on scarcely developed soils. It forms
with or without white adaxial faces; leaves some- open woodland alone or mixed with Juniperus spp.,
times producing a few resin drops. Stomata: leaves Pinus nelsonii, P. pinceana, Quercus, Yucca, Agave,
amphistomatic, or epistomatic in one variety, in Cactaceae (e.g. Opuntia), Arctostaphylos, Ceanothus,
23(4) lines on the convex abaxial face (or none), Arbutus and other shrubs of dry, hot areas; at higher
670 in 23(4) lines on each adaxial face. Pollen cones and/or moister sites it forms part of a mixed pine-
small, ca. 5 3mm, yellowish. Seed cones solitary, oak woodland or forest including, e.g. Pinus arizo-
paired or more rarely in whorls of 3, on very short, nica, P. engelmannii, P. leiophylla var. chihuahuana,
35(8)mm long peduncles remaining with fallen and P. pseudostrobus in the southeastern part of its
cones. Mature cones seemingly sessile, irregularly range. The climate is warm and dry, with annual
globose or ovoid-globose when closed, spreading precipitation varying from 380650mm and a dry
often wider than long when opened, with a flat- season of 78months. Frost may occur at higher
tened base, irregular in size and shape, often resin- elevations in the interior, but is infrequent. There is
ous, (2)35(7.5) 36(7)cm when open. Seed an important mutualist relationship with the corvid
scales parting easily and widely, more or less weakly birds Aphelocoma coerulescens and Gymnorhinus
attached to the rachis and hence moveable, up to cyanocephalus (Tomback & Linhart, 1990), which
1520mm wide, concavo-convex, with 12 deep feed on the seeds and cache them, thereby providing
seed cavities. Apophysis raised, transversely keeled an effective dispersal mechanism.
or radially keeled, rhombic to pentagonal in out-
line but irregular, in all shades from yellowish green Uses
or ochraceous to reddish brown, sometimes lus-
trous. Umbo dorsal, flat or raised and curved, with Although not a timber tree in most areas due to
a minute prickle. Seeds obovoid-oblique, 1016 its low stature and low and heavy branching, Pinus
610mm, greyish brown to blackish grey, or light cembroides is nevertheless an economically impor-
brown; integument thick, 0.51(1.1)mm; mega- tant species of pine in Mexico. Its principal value
gametophyte (endosperm) pinkish or white when for local economies lies in the edible seeds (pio-
fresh. Seed wings absent. nes), which are regularly harvested and marketed.
Further use is made of its wood for carpentry, or
Distribution sometimes for timber where there is no other pine
species available, as in Baja California Sur. Due to its
SW USA: SE Arizona, SW New Mexico, SW Texas; adaptation to semi-arid environments and extensive
Mexico: from the Sierra Madres to South-Central range it is also an important shrub or tree for aspects
Mexico and in Baja California (Sierra de la Laguna). of land management, such as watershed protection,
TDWG codes: 76 ARI 77 NWM TEX 79 MXC-DF prevention of erosion, and as a shade tree in agro-
MXC-ME MXC-PU MXC-TL MXE-AG MXE-CO forestry. Pinyon pines are uncommon in cultivation
MXE-CU MXE-DU MXE-GU MXE-HI MXE-NL and mostly seen in arboreta and botanic gardens and
MXE-QU MXE-SL MXE-TA MXE-ZA MXG-VC in some urban landscaping schemes mainly in SW
MXN-BS MXN-SO MXS-JA USA
by short sheaths, persisting 48 years, spreading, in California it is a component of the mixed coni-
(2)37(8)cm long, rigid, usually curved and fer forest as well as subalpine conifer woodland and
twisted, 0.72(3)mm wide; margins minutely ser- meadows with numerous conifer species. Here soils
rulate; leaf colour light yellow-green to dark green; are more nutrient rich and fires are less frequent, so
apex obtuse, acute or acuminate; stomata in lines Pinus contorta does not attain dominance. As a com-
on all surfaces. Pollen cones in small clusters, spi- ponent of the mixed conifer forest it can attain 50m
rally arranged, ellipsoid to cylindric, 515mm long, in height, with 1m d.b.h., and live to a considerable
orange-red turning red-brown. Seed cones solitary age. In other areas, such as large tracts of Yellowstone
or in whorls of 25, nearly sessile to short pedun- Park in Wyoming, Lodgepole pine appears to be
culate, spreading or reflexed, variably persistent self-perpetuating as the only tree species capable of
674 and variably serotinous, sometimes remaining on growing in a more dynamic environment character-
branches and closed for many years, ovoid but often ized by frequent fires.
with an asymmetrical base, (2)35(6)cm long,
broadly ovoid to nearly globose when open. Seed Uses
scales thin woody, rigid, brown. Apophysis more or
less rhombic in outline, variously raised, sometimes Lodgepole pine is a major timber tree in western
gibbous on the sun-exposed side near base of cone, North America. Its geographical varieties differ apart
transversely keeled, with a central, depressed, tri- from minor botanical characters also in growth per-
angular umbo armed with a variable, thin or blunt formance and maximum size, which is at least in
prickle up to 6mm long. Seeds obovoid, slightly flat- part due to environmental factors. As other coni-
tened, ca. 5mm long, red-brown with blackish spots fers, it attains greatest size in Oregon and California
or nearly black; wing 1012mm long. (var. murrayana) with trees over 50m tall and 2m
diam. In the interior, the tall, thin stems of densely
Distribution grown pines provided the lodgepoles, i.e. tent poles
for the bison-hide covered conical tents of the Plains
Western North America, from Yukon to Baja Indians with such famous tribes or nations as the
California and S Colorado. Dakota (Sioux) and Blackfeet. Today Lodgepole
TDWG codes: 70 ASK NWT-MK YUK 71 ABT BRC 73 pine is put to all traditional (European) uses com-
74 SDA 76 79 MXN-BC mon to pine wood, but mass production is for the
pulp industry or increasingly the manufacture of
Ecology so-called structural particleboard, where chips are
glued into boards for interior construction. Pinus
Pinus contorta occupies a large part of the North contorta is sometimes planted as a shelter tree on
American West with in particular a vast latitudinal barren sites, but otherwise uncommon in cultiva-
range. It consequently has a wide ecological ampli- tion; only a limited number of cultivars are known.
tude and grows form near sea level to 3350m or per- Hybrids between P. contorta and P. banksiana have
haps higher and from the relatively mild but cool and been generated by foresters in the USA in attempts
rainy Pacific coast to the cold and continental inte- to produce trees suitable for plantations.
rior of the northern Rocky Mountains. Precipitation
consequently ranges from only 250mm at low eleva- 3 varieties are recognized:
tions in the interior to 5000mm along the northern
coast. In the interior, Lodgepole pine forms pioneer
stands of great density after forest fires and can Pinus contorta Douglas ex Loudon var. contorta.
form monotypic stands of great extent, especially Type: USA: Washington, D. Douglas s.n.
on infertile soils. In other sites it is associated with (lectotype K).
many western conifers, most commonly in the north
with Picea glauca and mixed with Betula papyrifera Description
or Populus tremula; at higher altitudes with Tsuga
mertensiana, Picea engelmannii and Abies lasiocarpa. Leaves 25cm long, rarely longer, slender, 0.71.2(
Further south, the species diversity increases and 1.5)mm wide. Seed cones solitary or in whorls of 25,
asymmetrical near base, persistent and variously Pinus contorta Douglas ex Loudon var. murrayana
serotinous; umbos armed with a slender prickle to (Balf.) Engelm., in Watson, Bot. California 2: 126.
6mm long. 1880. Pinus murrayana Balf., in Murray, Bot.
Exped. Oregon 8: 2. 1853; Pinus contorta Douglas
Distribution ex Loudon subsp. murrayana (Balf.) Engelm.,
Trans. St. Louis Acad. Sci. 4: 177. 1880. Type: USA:
NW coast of North America, from S Alaska to N California, Siskiyou Mts, J. Jeffrey 740 (holotype E).
California. Fig. 218
TDWG codes: 70 ASK 71 BRC 73 ORE WAS 76 CAL
Description
Conservation 675
Trees to 50m tall; trunk to 12m d.b.h. Leaves
IUCN: LC 58cm long, 12mm wide. Seed cones solitary or
in pairs, mostly symmetrical or nearly so, opening
soon and falling soon after seed dispersal; apohyses
Pinus contorta Douglas ex Loudon var. latifolia transversely keeled but not gibbous; umbo with a
Engelm., in Watson, Botany (Fortieth Parallel): 331. small prickle.
1871. Pinus contorta Douglas ex Loudon subsp. lati-
folia (Engelm.) Critchf., Publ. Maria Moors Cabot Distribution
Found. Bot. Res. 3: 107. 1957; Pinus divaricata
(Aiton) Dum.-Cours. var. latifolia (Engelm.) Boivin, USA: California, Nevada, Oregon, S Washington;
Naturaliste Canad. 93: 272. 1966. Type: USA: Utah, NW Mexico: Baja California Norte.
Uintah Mts., S. Watson [Clarence Kings Exped. TDWG codes: 73 ORE WAS 76 CAL NEV 79 MXN-BC
40th Parallel] 1110 (holotype A).
Conservation
Description
IUCN: LC
Leaves (4)58cm long, 12(3)mm wide. Seed
cones solitary or in pairs, 46cm long, strongly
asymmetric and recurved on branches, persis- Pinus coulteri D. Don, Trans. Linn. Soc. London 17:
tent and variously serotinous; apophyses near base 440. 1836. Pinus ponderosa Douglas ex C. Lawson
of cone often gibbous; umbo with a small, blunt subsp. coulteri (D. Don) E. Murray, Kalmia 12: 23.
prickle. 1982. Type: USA: California, Monterey Co., Santa
Lucia Mts., Cone Peak, T. Coulter s.n. (holotype
Distribution TCD). Fig. 219, 220
Description
Etymology Distribution
The species epithet cubensis refers to Cuba, where it E Cuba: From the Sierra Maestra and the Sierra de
is endemic. Nipe E into the highlands terminating the eastern
part of the island.
Vernacular names TDWG codes: 81 CUB
Description Ecology
Shrubs 15m tall, diam. of stems to 1525cm; mul- The altitudinal range of P. culminicola is 3000
tistemmed, or very low branched, individuals form- 3700m a.s.l., which includes the highest ridges of
ing a spreading, in places dense vegetation. Bark these mountains. Its habit is very similar to other
thin, scaly, exfoliating in small, irregular plates, mountain dwarf pines, e.g. P. mugo in Europe and
grey-brown, soon weathering grey. Branches often P. pumila in NE Asia. Adaptation to blasting, ice-
prostrate to assurgent, the higher order branches or sand-laden winds and a short growing season is
assurgent to erect. Shoots short, thick, light brown responsible for this habit. Soils are mostly rocky and
turning grey. Cataphylls small, narrowly triangu- calcareous. Climatic conditions are not well known
678 lar to subulate; apex caudate; margins erose, light due to a lack of weather stations at these summits, but
brown, weathering blackish grey. Vegetative buds precipitation, some of it as snow, is probably abun-
broadly ovoid; terminal bud 610mm long; lateral dant. On Cerro Potos, the species forms extensive
buds smaller, slightly resinous. Fascicle sheaths ini- monocultures of close-packed individuals. It occurs
tially 68mm long, in mature fascicles the scales there with scattered, stunted P. hartwegii, which indi-
separate and recoil, forming a rosette at the base cates that the climatic tree line is not reached there
of fascicle, straw-coloured to grey, semi-persistent at around 3700m. Somewhat lower, on the Sierra La
but mainly falling before the leaf fascicles. Leaves Marta, Coahuila, P. culminicola has been found in a
in fascicles of 5 (very rarely 46), erect to assurgent, scrub-community with Quercus spp., Arctostaphylos,
persisting 23 years, curved, rigid, 35cm long, Ceanothus, Agave and grasses; on the Cerro La
0.91.3mm wide, very remotely serrulate to entire, Viega and the Sierra de Arteaga, Coahuila, a simi-
obtuse, greyish green, whitish or glaucous adaxi- lar vegetation, but also with Abies and Pseudotsuga,
ally. Stomata restricted to both adaxial faces, with are reported growing scattered with P. culminicula.
45 lines on each face. Pollen cones ovoid-oblong Pollen dispersal has been reported on Cerro Potos
when mature, 58mm long, yellowish, turning yel- to occur in late July, at 3690m, which indicates a late
lowish brown. Seed cones subterminal, solitary or fertilization and short growing season.
in pairs on short, stout peduncles which are covered
with subulate-caudate cataphylls. Mature cones sub- Conservation
globose when still closed, opening with a flattened
base and remote, spreading fertile seed scales, then This species is vulnerable to fire during long dry peri-
34.5 35cm. Seed scales parting and spreading ods. In recent years devastating fires have destroyed
wide except the smaller, infertile proximal scales, large parts of the population on Cerro Potos and
irregular, often curved, thin, concavo-convex, with regeneration, if it occurs, will be very slow.
12 deep, cup-like depressions holding the seeds. IUCN: EN [B1ab (iiv), B2ab (iiv)]
Apophysis slightly raised, transversely keeled,
rhomboid to pentagonal in outline, radially rugose Uses
when dry; margins crenate or angular, colour yel-
lowish brown, often resinous. Umbo dorsal, slightly This species is not used commercially, although
raised, rhombic in outline, with or without a min- locally it may be used for firewood. It should be
ute prickle. Seeds obliquely obovoid, 57 45mm, suitable as a low bushy pine for large rockeries, has
with a 1mm thick integument, brown. Seed wings attractive foliage, and deserves to be taken into cul-
absent on seeds parted from the scales. tivation for horticulture. It has established itself well
in the rock garden of the Royal Botanic Garden in
Distribution Edinburgh, Scotland, from seed collected by Michael
Frankis in 1991 and another, slightly later gathering.
Mexico: Coahuila, Nuevo Len. It remains very rare in cultivation; it seems that the
TDWG codes: 79 MXE-CO MXE-NL trade is often very slow in recognizing potentially
good, albeit perhaps slow growing, conifers for the
garden.
Pinus dalatensis Ferr, Bull. Soc. Hist. Nat. long and 10mm wide, the latter often with dark
Toulouse 95: 178. 1960. Pl. 28 longitudinal streaks.
This species is named after Dalat in Viet Nam, a city Pinus dalatensis is considered by Vietnamese bota-
near which it was discovered. nist to be a variable species (Nguyen Tien Hiep et
al., 2004). This species was studied in the field by
Vernacular names Businsk (1999, 2004), who not only discovered sev-
eral new populations in formerly unknown locali-
Dalat pine; Thng Lt (Vietnamese). ties, but described them under two subspecies, one 679
with a new variety. The discriminating characters
Description given in his papers are largely continuous and over-
lapping, but it seems probable that some of the vari-
Trees to 30(40)m tall; trunk to 2(2.5)m d.b.h., ation is sufficiently consistent within the disjunct
straight and columnar. Bark smooth in young populations to merit recognition as varieties. I have
trees and on branches, becoming rough and scaly retained them here, more or less provisionally, under
on trunk, breaking into small, grey-brown plates. the ranking given by Businsk, and listed only those
Branches wide spreading, self-pruning, forming few chracter states that appear to more or less sepa-
domed or umbrella-shaped, broad crowns in older rate them. It has to be emphasized (again) that geo-
trees. Foliage branches slender, smooth, new shoots graphical disjunction is not a taxonomic character.
glabrous or pubescent, often glaucous or pruinose, Some genetic research seems appropriate here.
becoming pale brown or reddish brown. Buds
small, conical or ovoid, acute, not resinous; cata- Distribution
phylls appressed, orange-brown. Leaves in fascicles
of 5, with deciduous basal sheaths, straight, spread- Viet Nam: Binh Tri Thin, Dac Lac, Gia Lai-Cng
ing, (3)510(14)cm long, slender, lax or pliant, Tum, Lm Dng; Lao PDR: Nakai Nam Theun NPA.
0.61(1.2)mm wide; margins mostly minutely ser- TDWG codes: 41 LAO VIE
rulate; leaf colour green or glaucous green (often a
combination, with abaxial face green); apex acute; Ecology
stomata in fine lines on the two adaxial faces. Pollen
cones in small clusters at base of new shoots, spirally Pinus dalatensis is a montane pine occurring at alti-
arranged, ovoid to short cylindrical, yellow. Seed tudes between 1400m and 2300m a.s.l. in tropical
cones solitary or in whorls of 23 on stout, 14cm evergreen forest. This species forms small stands
long, scaly peduncles, persisting some time after of a few to about 30 trees surrounded by evergreen
seed dispersal, initially erect but pendulous when angiosperm forest, dominated by members of the
grown to full size, extremely variable in size, 623cm Fagaceae and Lauraceae. In most localities, the pines
long, straight or curved ellipsoidal or cylindrical, occupy rocky outcrops or steep ridges and adjacent
opening to 59cm wide, resinous, green ripening to slopes where competition from broad-leaved trees is
dull or lustrous brown. Seed scales few or numer- less intense. In more favourable sites on flat moun-
ous depending on cone size, flat or boat-shaped, thin tain ridges or in foothills near streams, often on yel-
woody, flexible; basal scales appressed, rarely some low ferralitic soils, it becomes a large, emergent tree.
recurved. Apophysis large, thin or slightly thick- Associated conifers found are Pinus krempfii, also an
ened, more or less rhombic or with rounded distal emergent, Fokienia hodginsii, and Dacrydium elatum
margin, straight or slightly incurved, striated or in the southernmost part of the Central Highlands. In
grooved towards a terminal, 410mm wide, sunken, most cases, this pine can grow rapidly up to reach the
flat or obtuse umbo. Seeds varying in size with canopy, so it is semi-shade tolerant, but seedlings are
the seed scales (especially the wings), with maxi- intolerant of shade, so some form of canopy distur-
mal lengths of 810mm and wings to 2030mm bance has to precede regeneration, e.g. periodic fires.
680
Plate 28. Pinus dalatensis. 1. Habit of tree. 2. Branchlet with leaves. 3. Leaves. 4. Seed cones, one green,
one ripe. 5. Seeds.
Uses
Pinus dalatensis Ferr subsp. procera Businsk,
A valuable timber tree, but very rare and logged Candollea 54: 133. 1999. Type: Viet Nam: Gia Lai
only incidentally. Its wood properties are similar to Prov., Ngoc Linh Mountains, Mt. Ngok Niay,
those of Pinus wallichiana, a native of the Himalayas, R. Businsk 44114 (holotype G).
and other so-called white pines. It is not known in
cultivation. Description
2 subspecies and 2 varieties are recognized: New shoots (mostly densely) pubescent. Seed cones
923cm long.
Distribution IUCN: NE
Note on conservation status: Although the species
Viet Nam: Binh Tri Thin, Dac Lac, Lm Dng (Chu as a whole has been assessed as VU, the status of
Yang Sinh Massif, Da Lat environs); Lao PDR: Nakai the separate infraspecific taxa, although logically
Nam Theun NPA. meeting the criteria for at least VU, has not been
TDWG codes: 41 LAO VIE evaluated.
Conservation
IUCN: NE
Pinus densata Mast., J. Linn. Soc., Bot. 37: 416. and to some extent (via E. H. Wilsons field obser-
1906. Pinus tabuliformis Carrire var. densata vations) with P. sylvestris. Both species occur far to
(Mast.) Rehd., J. Arnold Arbor. 7: 23. 1926. Type: the NE from the known range of P. densata. The
China: Sichuan, Yalong River, E. H. Wilson 3015 hybrid status of this species has later been inferred
(holotype BM). from morphological intermediacy of character
states in comparison with P. yunnanensis, with
Etymology which it is partially sympatric, and P. tabuliformis,
which occurs also further north in China. Some
The species epithet is perhaps from Latin densatio molecular evidence (DNA sequence data) supports
= thickening (condensation) and may refer to the this hypothesis, possibly with the involvement of an
682 apophyses; if so it is not used in its original sense. unknown, third species from the Tertiary. Due to
ecological as well as geographical separation, there is
Vernacular names no indication that these species form new hybrids at
present in their natural habitats. It seems to me that
Gaoshan pine; gao shan song (Chinese) it is appropriate to treat this taxon not as a nothospe-
cies but as a species. Its hybrid status has not been
Description demonstrated beyond doubt, and given that hybrid-
ization is probably a major cause of speciation in
Trees to 30m tall; trunk to 1.3m d.b.h. Bark scaly, plants generally, we would not gain much relevant
forming irregular plates and longitudinal fissures, information if we did prove it.
dark grey-brown. Branches spreading, forming a
broadly domed or open crown in older trees. Foliage Distribution
branches stout, new shoots glabrous, rough with
pulvini from fallen leaf fascicles, lustrous yellowish China: S Qinghai, W Sichuan, NW Yunnan, E Xizang
brown, turning reddish brown. Buds ovoid-conical, [Tibet].
acute, the terminal bud ca. 15mm long, slightly res- TDWG codes: 36 CHC-SC CHC-YN CHQ CHT
inous; cataphylls with white fringed margins, acumi-
nate. Leaves in fascicles of 2(3), held by a persistent, Ecology
1015mm long but shortening basal sheath, straight,
814cm long, rigid, slightly twisted, 11.5mm wide, Pinus densata occurs in high mountains at altitudes
dull green; margins minutely serrulate; apex acute; from 2600m to 4000m a.s.l. or even above this line
stomata in fine lines on all faces. Pollen cones in to 4200m. It forms open pure stands at the highest
clusters at base of new shoots, spirally arranged, elevations, but becomes mixed with other pines, e.g.
short cylindrical, ca. 2cm long, yellow. Seed cones P. armandii and P. yunnanensis, below 3000m a.s.l.
solitary or in pairs, sessile or very short pedunculate, It is presumed that its hybrid characters include a
persistent but opening widely, spreading or curved greater tolerance to frost than either of its putative
down, narrowly ovoid when closed, 46cm long, parents, P. tabuliformis and P. yunnanensis, enabling
when opened 47cm wide. Seed scales oblong, thin it to colonize right up to the present tree line in west-
woody, rigid, brown; apophyses prominently raised, ern Chinas great mountain system.
more or less rhombic, strongly transversely keeled,
lustrous chocolate brown; umbo dorsal and central, Conservation
armed with a short prickle. Seeds ovoid to ellipsoid,
46mm long, light brown to grey-brown; wing IUCN: LC
1520mm long.
Uses
Taxonomic notes
There is presently little economic use of this spe-
Maxwell Masters (op. cit.) described this pine as a cies, primarily because it occurs in high mountains
new species and compared it with Pinus densiflora in often inaccessible places. A traditional use at least
and indeed the species itself, have made their way cal, 2040mm long, pink-purplish, turning light
to Europe, probably because there P. sylvestris offers brown. Seed cones subterminal, solitary or in pairs
similar opportunities for horticultural experimen- or whorls of 34. Mature cones on short, persistent
tation. The Japanese horticultural art bonsai also peduncles, leaving a few scales on the branch when
makes use of this species. Needles and extracts from falling, variable in size and shape, typically ovoid-
them are used in traditional medicine such as aro- oblong, with an oblique base, or curvate to cornute
matherapy. The pollen is edible and also used as (often by insect-damage), 1535 815cm when
medicine. open. Seed scales parting when the seeds are mature,
spreading wide, straight or slightly recurved, thick
woody, rigid or flexible with some force. Apophyses
Pinus devoniana Lindl., Edwardss Bot. Reg. 25: mostly raised, transversely keeled, rhombic in outline, 685
62. Aug 1839. [Allg. Gartenzeitung 7: 324. 1839] with irregular sides, up to 25mm wide, often radially
Type: Mexico: Hidalgo, Mineral del Monte, Cerro striated, in various shades of brown. Umbo dorsal,
Ocotillo, C. T. Hartweg s.n. (lectotype W). Fig. 222 raised, flat or depressed, terminating in a small, usu-
ally deciduous prickle, grey-brown. Seeds obliquely
Pinus michoacana Martnez, Anales Inst. Biol. Univ. broad ovoid, flattened, 810 57mm, light brown,
Nac. Mxico 15: 1. 1944. often with dark spots. Seed wings obliquely ovate to
oblong, with a straight side, 2535 1015mm, light
Etymology brown with darker stripes. Many seedlings develop-
ing a grass stage with delayed apical growth of stem.
This species was named in honour of William
Spencer Cavendish, sixth Duke of Devonshire Taxonomic notes
(17901858), who had a great interest in gardening.
This species is still known in many books on coni-
Vernacular names fers by its name given by Maximino Martnez, Pinus
michoacana. Martnez (1945, 1948), who worked
Michoacan pine; pino blanco, pino lacio, ocote on the pines of Mexico during the difficult years of
gretado (Mexico) World War II, considered it likely that his new spe-
cies would correspond with some of John Lindleys
Description names, among which P. devoniana was mentioned,
but at the time he had no access to these original
Trees to 2030m tall, d.b.h. to 0.81m. Trunk publications. The complexity of forms and many of
monopodial, usually erect; bark thick, scaly, very the names that were given to these have been united
rough, with elongated plates divided by deep longi- by Farjon & Styles (1997) in a comprehensive mono-
tudinal black fissures, reddish brown to dark brown. graph on Latin American pines under the earliest
Branches spreading and assurging, or ascending available name Pinus devoniana Lindley. Hopefully,
near the top, forming an open, broad pyramidal or the results of thorough taxonomic work published
domed crown. Shoots 1520mm thick, rigid, curved, in specialized literature that strictly follows the
very rough and scaly. Cataphylls up to 2025mm rules of botanical nomenclature will eventually filter
long, subulate, scarious, recurved or reflexed, dark through to become established in the literature aim-
brown to blackish grey, with erose-ciliate margins. ing at a more general audience.
Vegetative buds large; terminal bud 2040mm long,
but lateral buds smaller, ovoid-acute, not resinous. Distribution
Fascicle sheaths to 3040mm long, often very resin-
ous. Leaves in fascicles of 5, rarely 4 or 6, persisting Mexico: Sinaloa, Nayarit, Jalisco, Zacatecas,
23 years, rigid, straight, or flexible and drooping, Aguascalientes, San Lus Potos, Quertaro, Hidalgo,
(17)2540(45)cm long, 1.11.6mm wide, with Michoacn, Mxico, Distrito Federal, Morelos,
serrulate margins, acute-pungent, lustrous green. Tlaxcala, Puebla, Veracruz, Guerrero, Oaxaca and
Stomata on all faces of leaves. Pollen cones cylindri- Chiapas; in Guatemala in the southern highlands.
TDWG codes: 79 MXC-DF MXC-ME MXC-MO but otherwise it would do fine in much of California,
MXC-PU MXC-TL MXE-AG MXE-HI MXE-QU the Mediterranean, Australia, and New Zealand, to
MXE-SL MXE-ZA MXG-VC MXN-SI MXS-GR mention just a few regions where gardening and tree
MXS-JA MXS-MI MXS-NA MXS-OA MXT-CI 80 GUA planting are a major enjoyment for many.
Ecology
Pinus douglasiana Martnez, Madroo 7: 4. 1943.
This species is a constituent of relatively open, often Type: Mexico: Nayarit, Jal, Cerro Juanacata, M.
secondary pine-oak forests, or it occurs with Pinus Martnez 3429 (holotype MEXU).
oocarpa invading burned mountainsides. It is well
686 adapted to withstand fires by its juvenile grass stage Etymology
similar to that of P. palustris of the SE USA. The
pines most commonly associated with it vary some- This species was named after the plant collector
what from north to south and include P. oocarpa, David Douglas (17981834), who, however, never
P. montezumae, P. pseudostrobus, and P. maximinoi visited Mexico.
and less frequently P. cembroides at lower altitudes
and P. hartwegii at higher altitudes. Common are Vernacular names
also Quercus, Liquidambar, and in the understorey
Calliandra, Leucaena, Acacia, Dodonaea, Gaultheria, Douglas pine; ocote, pino hayarn (Mexico)
and Mimosa. Its altitudinal range is (700)900
2500(3000)m a.s.l. Pinus devoniana grows on a Description
variety of soils, often of volcanic origin. The climate
is warm-temperate to subtropical, with annual pre- Trees to 2045m tall, d.b.h. to 0.81m. Trunk
cipitation 10001500mm and a dry season from monopodial, straight; bark rough, scaly, divided into
November to May. large, irregular plates and deep fissures, dark red-
dish brown, weathering grey-brown. Branches long,
Conservation slender, spreading, or ascending in upper part of
crown. Shoots uni-nodal, dark brown, not glaucous.
IUCN: LC Cataphylls subulate-caudate, spreading or recurved,
with erose-ciliate margins, brown. Vegetative buds
Uses ovoid-conical; terminal bud 1525 1015mm, but
lateral buds smaller, not resinous. Fascicle sheaths
Pinus devoniana is a common tree throughout the persistent, remaining long, (15)2035mm, weath-
mountainous parts of S Mexico and Guatemala, ering greyish brown. Leaves in fascicles of 5, rarely 4
often encountered in open, degraded pine-oak or or 6, persisting 22.5 years, slender, lax or sometimes
pine forest. As it is not a very tall, straight-boled more rigid, 2235cm long, 0.71.2mm wide; mar-
tree and has branches low on the trunk, it is not gins serrulate; apex acute; leaf colour light yellow-
often selected as a timber tree. On the other hand, ish green to glaucous green. Stomata on all faces of
it is locally used and, like other pines, increasingly leaves. Pollen cones numerous, forming long, spicu-
for firewood by the growing rural population. Uses late clusters, cylindrical, 2025 45mm, pinkish
of timber are put to fence posts, boxes, furniture, brown at maturity. Seed cones solitary, in pairs or
toll handles, and other woodware and, mixed with whorls of 34 on stout, recurved, 1015mm long
other pine wood, wood chips glued and compressed peduncles which fall with the cone. Mature cones
to particleboard. It is virtually unknown in hor- ovoid to ovoid-attenuate when closed, often slightly
ticulture despite the fact that it is one of the most curved, more broadly ovoid with a flattened base
strikingly beautiful pines, with perhaps the longest when opened, then 710 57cm. Seed scales part-
needles in the entire genus and large cones and a ing to release the seeds except at base, (thin) woody,
deep red-brown bark. Provenance is crucial to estab- oblong, straight or slightly curved. Apophyses nearly
lish whether some trees could turn out to be hardy, flat or raised and transversely keeled, radially striate
or grooved, rhombic or pentagonal in outline, nearly sought out for tree-breeding purposes, perhaps due
symmetrical around cone, 1117mm wide, light to the fact that it is relatively unknown to foresters.
brown, reddish brown or dark brown. Umbo dorsal, It is perhaps present in a few arboreta or pineta in
raised, transversely rhombic, darker than the apoph- regions with a mild climate, but not generally cul-
ysis, prickle absent. Seeds obliquely obovoid, slightly tivated for horticulture. It has been tried in forestry
flattened, 45 33.5mm, light grey or brown, often plantations, but is not in common use.
with dark spots. Seed wings oblong, with a straight
and a curved side, 1824 79mm, light yellowish
brown, translucent, sometimes with a darker brown Pinus durangensis Martnez, Anales Inst. Biol.
tinge. Univ. Nac. Mxico 13: 23, f. 14. 1942. Type:
Mexico: Durango, El Salto, Las Adjuntas, M. 687
Distribution Martnez 3477 (holotype MEXU). Fig. 223
Mexico: mainly in Jalisco, Michoacn, Mxico and Pinus martinezii E. Larsen, Madroo 17: 217.
N Morelos, but extending northward into Nayarit 1964; Pinus douglasiana Martnez var. martinezii
and the crest of the Sierra Madre Occidental on the (E. Larsen) Silba, Phytologia 68: 49. 1990.
border line of Sinaloa and Durango, southwards it
occurs locally in Guerrero and Oaxaca. Etymology
TDWG codes: 79 MXC-ME MXC-MO MXC-PU
MXE-DU MXN-SI MXS-GR MXS-JA MXS-MI The species epithet durangensis refers to the Mexican
MXS-NA MXS-OA State of Durango, from where it was first described.
This species is a constituent of mostly mixed pine Durango pine; ocote, pino blanco, pino real (Mexico)
or pine-oak forests at elevations ranging from
(1100)14002500(2700)m a.s.l. in warm to tem- Description
perate climatic zones. The annual precipitation dif-
fers with altitude but is roughly around 1000mm Trees to 3540m tall, d.b.h. to 0.81m; trunk mono-
in most areas. Common associated pines are Pinus podial, erect, straight. Bark rough, scaly, breaking
pseudostrobus, P. herrerae, P. leiophylla, P. lawsonii, into large, irregular, elongated plates divided by lon-
P. ayacahuite in the southern part of its range, some- gitudinal, shallow fissures, dark brown, weathering
times P. oocarpa and P. devoniana at lower elevation grey. Branches long, slender, with lower branches
and drier sites respectively. At the highest and wet- usually curved downward, ultimately more or less
test sites P. douglasiana can occur with Abies, Picea pendant. Shoots uni-nodal, orange-brown or red-
(in Durango) or Cupressus lusitanica. In many areas dish brown, usually glaucous. Cataphylls ca. 15mm
Quercus spp. are codominant; a shift in forest com- long, 34mm wide at base, subulate, soon reflexed,
position towards broad-leaved trees may occur also scarious, with erose-ciliate margins and caudate
by selective cutting of pines. apex, dark brown. Vegetative buds ovoid; termi-
nal bud 1520(25) 1015mm, but lateral buds
Conservation smaller, not resinous. Fascicle sheaths 2030mm
long, persistent but reduced to 1015mm. Leaves
IUCN: LC in fascicles of (4)56(7, rarely 8), persisting 22.5
(3?) years, straight or slightly curved, 1424cm
Uses long, 0.71.1mm wide; margins serrulate, acute to
pungent; leaf colour yellowish green to glaucous
Pinus douglasiana is, along with other species with green. Stomata on all faces of leaves. Pollen cones
which it often occurs, an important timber tree ovoid-oblong to cylindrical, yellowish brown when
in most of its range. It is, however, not especially mature. Seed cones subterminal, solitary, in pairs
Ecology Etymology
In the Sierra Madre this species is an important con- The species epithet means spiny or prickly, but it is
stituent of the yellow pine forest, where it occurs not clear what Philip Miller found so spiny in this pine.
in pure stands or mixed with several other species
of pine, e.g. P. arizonica, P. leiophylla, and P. engel- Vernacular names
mannii, or in pine-oak forests. Its altitudinal range
is extensive: (1400)16002800(3000?)m, but it is Shortleaf pine
most common at 20002800m a.s.l. Climatically,
the Sierra Madre is warm-temperate, but with cold Description
spells during the short winter at the higher eleva-
tions. Annual precipitation varies between 700 Trees to 3540m tall; trunk to 1.6m d.b.h. Bark
1200mm, most of which occurs in the summer. This breaking into irregularly rectangular or square,
pine is adapted to grow on shallow, rocky soils, but scaly plates, dark brown, slightly resinous. Branches
its better stands are found on deeper soils, where spreading, sometimes heavy but more slender and
it can successfully compete with most other pines. shorter in trees growing in closed canopy stands.
The soils are mostly derived from volcanic rock. Foliage branches slender, often more or less pen-
At the highest elevation P. durangensis occurs with dulous, new shoots purplish green, often glaucous,
Abies and/or Cupressus lusitanica, at the lowest with turning red-brown to grey, rough with persistent
Juniperus deppeana and Pinus oocarpa. Other pines pulvini after leaf fascicles have fallen. Buds ovoid
are P. montezumae, P. teocote and in the southern to ovoid-cylindric, 510mm long, strongly resin-
part of its range P. ayacahuite can occur with it. ous. Leaves in fascicles of 2 or 3, held in a 1015mm
long fascicle sheath, persisting 35 years, spreading, land types. It can also be associated with P. taeda,
(5)611(12)cm long, straight or slightly curved which has a very similar distribution, but at least in
and flexible, ca. 1mm wide, yellowish green, grey- parts of its range occurs in a wetter habitat.
ish green or dark green; margins minutely serrulate;
apex short acute; stomata in fine lines on all surfaces. Conservation
Pollen cones in small clusters, spirally arranged,
cylindrical, 1.52cm long, pale purplish green to IUCN: LC
yellow, turning light brown. Seed cones solitary or
in whorls of 24, short pedunculate or nearly ses- Uses
sile, semi-persistent but opening soon, spreading,
47cm long, narrowly ovoid or ovoid-oblong when Shortleaf pine is an important commercial conifer 689
closed, broadly ovoid or ovoid-conical with a nearly species in the SE United States and both natural
flat base when open. Seed scales thin woody, rigid, stands and plantations are exploited for timber. The
uniformily dull brown below the apophysis without wood is of excellent quality, with orange or yellow-
a dark band; apophysis slightly raised, sharply keeled ish brown heartwood and creamy yellow sapwood;
transversally; umbo dorsal and central, armed with it is used for railway sleepers, construction lumber,
a short, stout prickle. Seeds ellipsoid, ca. 6mm long, indoor finishing like panelling, plywood, furniture,
grey mottled black, with a similarly patterned wing and kraft pulp and dissolving pulp; the latter prod-
1215mm long. uct feeds the paper industry. Most of the planta-
tion timber goes to pulping. There is a limited use
Distribution for amenity planting and in urban areas this pine is
planted to screen off residential areas from motor-
E and SE USA, from New York to E Texas across ways (major highways) or industrial areas. This
22 states. species has little significance in horticulture and is
TDWG codes: 74 ILL MSO OKL 75 CNT NWJ NWY rarely planted in gardens.
OHI PEN WVA 77 TEX 78 ALA ARK DEL FLA GEO
KTY LOU MRY MSI NCA SCA TEN VRG WDC
Pinus edulis Engelm., in Wislizenus, Mem. Tour N.
Ecology Mexico: 88. 1848. Pinus cembroides Zucc. var. edulis
(Engelm.) Voss, Mitt. Deutsch. Dendrol. Ges. 1907
Pinus echinata is a lowland pine with an extensive (16): 95. 1907; Pinus cembroides Zucc. subsp. edulis
range across the SE United States, mostly growing (Engelm.) E. Murray, Kalmia 12: 22. 1982. Type not
from ca. 150m up to the foothills of the Appalachian designated.
Mountains at ca. 600m a.s.l. It is absent in the
Mississippi Valley and its delta as well as in a narrow Etymology
to fairly wide coastal strip along the Gulf of Mexico
and Atlantic Ocean, and it does not extend into most The species epithet means edible and refers to the
of Florida. This indicates that it is primarily limited seeds.
by climatic factors, such as the 10 C average annual
temperature isoline at its northern limit and an aver- Vernacular names
age annual precipitation above 1000mm distributed
more or less evenly over a year as its southern limit. Pinyon pine, Two-needle pinyon pine, Colorado
It grows on a great variety of soils, but most have pinyon
a capacity for moist retention with a sandy loam or
silty loam texture and good drainage. Although this Description
species can form nearly pure stands, in most sites
it will be succeeded by broadleaved trees especially Shrubs or trees to 15(21)m tall, d.b.h. to 60cm.
oaks (Quercus spp.) except where thin soil overlies Trunk monopodial, short to medium size, strongly
rock. There P. echinata forms a minor component tapering, branching low. Bark thick, rough and scaly,
of various angiosperm-dominated forest and wood- breaking into small, irregular plates, divided by
irregular, shallow fissures, not easily exfoliating, red- States of the USA. It forms extensive, open stands
brown weathering grey-brown. Branches spreading commonly with one or more species of Juniperus
or ascending, irregularly disposed in most trees. known as Pinyon-Juniper woodland, which is one of
Crown broad conical or rounded, dense. Shoots the most widespread semi-arid vegetation types in
pale red-brown or tan, rarely slightly glaucous at North America. Summers are hot and winters cold,
first, soon grey, glabrous or minutely puberulent. but climatic conditions are varying with altitude and
Vegetative buds ovoid-oblong to ellipsoid; terminal latitude. Soils are commonly thin to sceletal or may
bud 510 35 mm, but lateral buds smaller, resinous, be absent altogether, with the trees growing from fis-
red-brown. Fascicle sheaths 57mm long, loosely sures in the sandstone, limestone, or shale. Recent
imbricate, soon recoiling, then pale straw-coloured sedimentation accumulates in the basins and valley
690 to grey, forming a small rosette at base of fascicle but bottoms, where grasses and sagebrush (Seriphidium
deciduous before the leaves fall. Leaves in fascicles of tridentatum) dominate, while at higher elevations
(1)2(3), spreading, persisting 46 years, upcurved, in the mountains the Pinyon-Juniper woodland
connivent, rigid, 24cm long, 0.91.5mm wide; gives way to open pine forest with Pinus ponderosa
margins entire or minutely serrulate; leaf colour vari- and Pseudotsuga menziesii. The altitudinal range of
able, dull green to glaucous green. Stomata: leaves P. edulis is 9103200m a.s.l. Juniperus monosperma
amphistomatic, with 24 lines on each face. Pollen and J. osteosperma are the most commonly associ-
cones in more or less elongated, spiculate clusters, ated junipers with P. edulis. The appearance is of a
ca. 7 3.5mm, yellowish to red-brown. Seed cones stunted forest as the free standing trees branch low
solitary, paired or more rarely in whorls of 3, on very and form wide spreading crowns while only attain-
short, 35mm long peduncles remaining with fallen ing modest height. Depending on openess, there
cones. Mature cones seemingly sessile, irregularly is an understorey dominated by shrubs of which
globose or ovoid-globose when closed, spreading Seriphidium (Artimisia) is most common and wide-
often wider than long when opened, with a flattened spread, supplemented by scrubby oaks (Quercus
base, irregular in size and shape, often resinous, spp.), Chrysothamus, Cercocarpus, Ephedra, Yucca,
35 36cm when open. Seed scales parting eas- and several others depending on geographical area,
ily and widely, narrowly and more or less weakly as well as grasses and other herbs.
attached to the rachis and hence moveable, con-
cavo-convex, with 12 deep seed cavities. Apophysis Conservation
prominently raised, transversely keeled or radially
keeled or ribbed, rhombic to pentagonal in outline Although the species is too widespread and abun-
but irregular, often angular or curved, variously dant to be threatened with extinction, the Pinyon-
brown. Umbo dorsal, flat or slightly raised, with a Juniper woodland as an ecosystem is under threat
minute prickle. Seeds obovoid or ellipsoid, 1015 in many places due to range improvement for the
69mm, greyish brown or light brown; integument grazing of cattle and sheep, causing the removal or
thick, 0.51(1.1)mm; megagametophyte (endo- degradation of the woodland over large areas. There
sperm) pinkish or white when fresh. Seed wings is growing awareness that these practices must stop,
absent when the seed is detached from the scale. as the habitat itself is increasingly found to be more
important than the production of more beef and
Distribution wool and much of the Pinyon-Juniper woodland
occurs on public lands.
SW USA: Arizona, S California, Colorado, New IUCN: LC
Mexico, W Oklahoma, NW Texas, Utah, S Wyoming.
TDWG codes: 73 COL WYO 74 OKL 76 ARI CAL UTA Uses
77 NWM TEX
Firewood from the Pinyon-Juniper woodland is the
Ecology most common use both past and present. Pinyon
pine wood has a higher than average heat value and
Pinus edulis is widely distributed in the interior basins, burns with a distinctive aroma (as does the juniper
plateaus, mesas and mountains of the Four Corner wood, which I prefer for my campfires when trav-
eling in the area). Large quantities were logged and 18)cm long, narrowly ovoid or ovoid-oblong when
its rough timbers used as props in the mining boom closed, broadly ovoid or ovoid-cylindric with a more
of the late 19th century. Poor growth form from the or less flattened base when open. Seed scales thin
foresters perspective renders the wood unsuitable woody, rigid, light brown proximally; apophysis lus-
for sawn timber, even though its quality matches trous brown, raised and sharply keeled transversally;
that of Ponderosa pine. The edible seeds are easy to umbo dorsal and depressed-pyramidal, armed with
harvest and in great demand as a delicacy; they may a short, stout prickle. Seeds ellipsoid, 67mm long,
constitute the economically most valuable product dark brown, with a wing 2030mm long. Seedlings
of the species. Crops can vary greatly from one year with or without a grass stage.
to the next and the slow growing pines do not per-
form well in plantations. Horticultural value is lim- Distribution 691
ited, although locally young trees are harvested as
Christmas trees. SE USA, from North Carolina to Mississippi, S to
Florida Keys.
TDWG codes: 78 ALA FLA GEO LOU MSI NCA SCA
Pinus elliottii Engelm., Trans. Acad. Sci. St. Louis 4:
186, t. 13. 1880. Ecology
Uses Conservation
Pinus fenzeliana Hand.-Mazz., Oesterr. Bot. Z. 80: length extremely variable at least between popula-
337. 1931. Pinus parviflora Siebold & Zucc. var. tions, 418cm long, leaves spreading or droop-
fenzeliana (Hand.-Mazz.) C. L. Wu, Acta Phytotax. ing, slender, flexible, the shorter leaves straight or
Sin. 5 (3): 143. 1956. Type: China: Hainan Island, curved, 11.5mm wide, glaucous green, with sto-
[?] Fenzel 55 (holotype WU). matal lines on the two adaxial surfaces; margins
minutely serrulate. Pollen cones in small clusters,
Pinus kwangtungensis Chun & Tsiang, Sunyatsenia short cylindrical. Seed cones variable in size and
7: 113. 1948; Pinus wangii Hu & W. C. Cheng var. shape, from short ovoid to long cylindrical, ini-
kwangtungensis (Chun & Tsiang) Silba, Phytologia tially erect on stout peduncles, becoming curved
68: 64. 1990; Pinus wangii Hu & W. C. Cheng subsp. down to pendulous, (3)515(17)cm long, solitary
694 kwangtungensis (Chun & Tsiang) Businsk, Acta or with 23 together. Seed scales soft woody, more
Pruhoniciana 68: 11. 1999. or less flexible at base, cuneate to oblong; apophy-
Pinus kwangtungensis Chun & Tsiang var. variifolia ses rhombic to oblong (at base and apex of cone),
Nan Li & Y. C. Zhong, Novon 7 (3): 262. 1997; Pinus curved or more or less straight, not recurved or
wangii Hu & W. C. Cheng subsp. variifolia (Nan Li more commonly recurved near cone base, becoming
& Y. C. Zhong) Businsk, Acta Pruhoniciana 68: 11. longitudinally furrowed in mature cones, ripening
1999. to yellowish brown or reddish brown, weathering
Pinus fenzeliana Hand.-Mazz. var. annamiensis grey-brown; apex thin or marginally thickened, usu-
Silba, J. Int. Conifer Preserv. Soc. 7 (1): 30. 2000. ally upcurved; umbo terminal, small and obtuse.
Pinus orthophylla Businsk, Willdenowia 34 (1): 229. Seeds obovoid or ellipsoid, 1015mm long, wingless
2004. when dispersed, or with a variously rudimentary to
Pinus eremitana Businsk, Willdenowia 34 (1): 234. small wing always shorter than the seed and easily
2004. detached.
This pine species has been named after Fenzel the Using a somewhat broader species circumscrip-
collector of the type specimen. tion here, I have united Pinus kwangtungensis with
P. fenzeliana, a name which must then be taken up
Vernacular names as the earliest one validly published for the broader
defined species. The variability of this species, with
hai nan wu zhen song, hua nan wu zhen song character states inconsistently distributed within
(Chinese names for two species here united into one). and among populations, has led to the naming of
many varieties and the classification of these with
Description more than one species. In Flora of China 4 (1999)
P. fenzeliana, P. kwangtungensis and P. wangii are kept
Trees to 50m tall, in many areas only to 2030m separate, the two former each with two varieties. In
tall; trunk to 1m d.b.h. Bark on young trees and this group of pines (section Quinquefolius, subsec-
branches smooth, thin, becoming scaly and flak- tion Strobus) the number of leaves per fascicle is five
ing, brown, dark brown or grey-brown on trunks and remarkably constant (very rarely a few fascicles
of larger trees. Branches spreading wide, forming with more than five, but never with fewer). The vari-
broad, umbrella-shaped or domed crowns. Foliage ety variifolia recently described by Nan Li and Y. C.
branches slender; young shoots pale brown, rarely Zhong (op. cit.) seems to be based on erroneous
glaucous, glabrous or rarely puberulent in grooves, observation. Recently, Businsk (2004) in a revi-
turning greyish brown. Buds ovoid to cylindrical, sion of Pinus subsection Strobus in Asia, described
slightly resinous; cataphylls dark brown. Leaves in two new species, one from Hainan Island in China
fascicles of 5, held by deciduous sheaths of flimsy, (P. orthophylla) and one from N Viet Nam
brown scales; some leaves may fall independently (P. eremitana) which both appear to belong here.
before the whole bundle falls off the twig, produc- The differences in character states tabulated in this
ing fascicles with seemingly fewer than 5 leaves. Leaf paper are not distinct, but have overlapping values,
Etymology Distribution
The species epithet refers to the very flexible W North America: Rocky Mountains from Alberta
branches. and British Columbia to New Mexico and Texas and
some isolated localities to the east, mountains of SE
Vernacular names California, Nevada, N Arizona, Utah; and extend-
ing into northern Mexico in scattered localities in
Limber pine, Rocky Mountain White pine Chihuahua, NE Sonora, Coahuila and Nuevo Len.
696
Plate 29. Pinus flexilis. 1. Habit of tree. 2. Branch with foliage. 3. Leaves. 4. Seed cone.
TDWG codes: 71 ABT BRC 73 COL IDA MNT WYO 74 Pinus flexilis E. James var. flexilis. Type: USA:
SDA 76 ARI CAL NEV UTA 77 NWM TEX 79 MXE-CO Colorado, El Paso Co., eastern flank of Pikes Peak,
MXE-CU MXE-NL MXN-SO 0.25miles north of Ruxton Park, Andresen J. W. &
[?] Barger A2125 (neotype SIU). Pl. 29
Ecology
Pinus flexilis E. James var. macrocarpa Engelm., in
Pinus flexilis is a conifer tree of the subalpine zone Rothrock, Rep. U.S. Geogr. Surv., Wheeler, 6, Bot.:
in the Rocky Mountains and the Basin and Range 258. 1879; Pinus novaemexicana P. Landry, Phytologia
region further west. In many respects its habitat 65: 479. 1989.
resembles that of Pinus albicaulis, which has a more
westerly and northerly distribution. Some isolated Description 697
populations, here recognized as var. reflexa, occur
on high peaks in northern Mexico. In the Black Hills Seed scales imbricate but spreading at maturity to
of North Dakota P. flexilis may grow as low as 900m release seeds; apophyses straight or slightly incurved,
a.s.l., in the southern Rocky Mountains it can occur not recurved. Seeds without a wing when detached
at 3800m. This species is one of several in the genus from the scale.
Pinus of western North America that can withstand
extreme conditions of climate on bare rock or scree Distribution
without any other vegetation cover. On these sites it
occurs either alone or with Pinus albicaulis and Abies W North America: Rocky Mountains from Alberta
lasiocarpa in the northern parts of its range and with to New Mexico, mountains of SE California, Nevada,
Pinus aristata in the SE and P. longaeva in the SW. At N Arizona, Utah, isolated localities in North Dakota,
lower altitudes it is usually only a minor component South Dakota and Nebraska.
of more diverse conifer forest in the south and with TDWG codes: 71 ABT BRC 73 COL IDA MNT WYO 74
Picea engelmannii, Pinus contorta, and Pseudotsuga NDA NEB SDA 76 ARI CAL NEV UTA 77 NWM
menziesii in the north. The seeds, which only have
rudimentary wings, are mostly dispersed by rodents Conservation
and birds, of the latter Clarks Nutcracker (Nucifraga
columbiana, Corvidae) is the most important vector. IUCN: LC
Uses
Pinus flexilis E. James var. reflexa Engelm., in
Although the wood of Limber pine is of high qual- Rothrock, Rep. U.S. Geogr. Surv., Wheeler, 6,
ity, as a timber tree the species is of minor impor- Bot.: 258. 1879. Pinus reflexa (Engelm.) Engelm.,
tance due to its occurrence at high altitudes, where Bot. Gaz. 7: 4. 1882; Pinus flexilis E. James subsp.
it is generally inaccessible and most of the trees do reflexa (Engelm.) E. Murray, Kalmia 12: 23. 1982.
not grow into straight boles. Planted trees have a Type: USA: Arizona, Pima Co., Santa Rita Mts.,
tendency to become multi-stemmed from a short E of Tucson, J. T. Rothrock 654 (lectotype MO).
base and grow rather slowly. Locally, it has been
used much for construction in pioneer times, but Pinus strobiformis Engelm. var. potosiensis Silba,
more recently its main uses have been restricted to Phytologia 68: 62. 1990.
carpentry and furniture, as well as firewood. The Pinus stylesii Frankis ex Businsk, Acta Pruhon. 88:
species has been introduced to Britain in 1851, but 6. 2008.
has remained rare in gardens there and elsewhere in
Europe. In the USA it is more common and several Seed scales imbricate, spreading when mature;
cultivars, most of which have glaucous blue needles apophyses slightly recurved. Seeds with a wing to
and more compact habits, are being grown and are 5mm or wing vestigial.
in the horticultural trade.
In a recent publication on the genus Pinus, Businsk Trees to 25m tall, but usually to 1518m; trunk to
(2008) separated the populations of white pines 0.81m d.b.h., usually branching low, sometimes
in the northern part of the Sierra Madre Oriental, multi-stemmed. Bark smooth, hard, irregularly exfo-
Mexico, as well as some outlier populations, as a dis- liating with large thin plates, exposing fresh patches,
tinct species P. stylesii. This concept includes P. strobi- which later turn from yellow-green or pale green to
formis var. potosiensis Silba and P. flexilis var. reflexa, greyish white and create a multi-coloured pattern on
both from Cerro Potos in Nuevo Len. Chris Earle branches and a grey-white trunk; bark on lower por-
(www.conifers.org) has considered that it may be tion of trunk can become rough and fissured. Main
698 of hybrid origin involving (originally) P. flexilis and branches long, spreading and ascending, forming a
P. strobiformis (= P. flexilis var. reflexa). As P. flexilis very wide, open crown. Foliage branches slender or
retreated from the area in post-glacial time, while stout, smooth, glabrous, yellowish green to olive-
P. strobiformis seems to have increased its range, green. Buds ovoid, reddish brown, not resinous.
fewer and fewer original hybrid trees remained Leaves in fascicles of 3, at first held by a basal sheath,
and the genes from P. strobiformis may eventu- the scales of which are shed in the second year,
ally subsume those from these earlier hybrid persistent 23 years, straight or curved and rigid,
trees. If that scenario is correct, the name P. flexi- spreading, 510cm long, slender, ca. 1mm wide,
lis var. reflexa would be the most appropriate for grey-green or dark green; stomata on the two adaxial
this taxon. faces. Pollen cones clustered at the beginning of new
shoots, spirally arranged, 7.513mm long, ovoid-
Distribution cylindrical. Seed cones solitary, lateral on shoot,
sessile or short pedunculate, robust, 1220(23)cm
USA: Arizona, New Mexico, SW Texas; Mexico: long, 811(13)cm wide when open, oblong-ovoid,
Chihuaha, Coahuila, Nuevo Len, NE Sonora. with a more or less level base, falling 23 years after
TDWG codes: 76 ARI 77 NWM TEX 79 MXE-CO maturity. Seed scales thick woody, rigid, 3.55cm
MXE-CU MXE-NL MXN-SO long and 22.5cm wide at mid-cone, spreading rela-
tively wide, with two deep cavities on adaxial surface
Conservation that hold the seeds; apophysis strongly developed,
reflexed, longitudinally ridged and grooved, apically
IUCN: NT strongly curved near base of cone; umbo dorsal,
with a recurved or hooked, obtuse apex. Seeds large,
2025mm long, ovoid-oblong to more or less cylin-
Pinus gerardiana Wall. ex D. Don, in Lambert, dric, asymmetrical, 812mm wide; wing rudimen-
Descr. Pinus, ed. 8, 2: p. s.n. inter 144 et 145, t. tary, remaining attached to the seed scale.
79. 1832. Type: India: Uttaranchal, Kumaon, Lt.
Gerard s.n. [Wallich Cat. No. 6064] (lectotype K-W, Distribution
here designated).
E Afghanistan; China, S Xizang (Tibet); India,
Etymology Jammu-Kashmir; N Pakistan.
TDWG codes: 34 AFG 36 CHT 40 PAK WHM-HP
This pine species was named after its European dis- WHM-JK
coverer Lt. Gerard, who brought it to the attention of
Nathaniel Wallich in Calcutta. Ecology
Liquidambar, Nyssa, Carya, Fagus, and Quercus and multi-nodal, smooth, reddish brown to grey-brown.
survives by overtopping them. This tolerance to Cataphylls lanceolate to subulate, straight, erose-cil-
shade is rather unusual for a pine and it is dimin- iate at margins, dark brown. Vegetative buds ovoid-
ished as it grows taller, so it often requires some conical to oval-oblong; terminal bud 1015mm long;
canopy opening to attain dominance. Unlike many lateral buds more ovoid and smaller, not resinous.
other pines it is susceptible to fire. It may occasion- Fascicle sheaths initially ca. 10mm long, persistent
ally grow with other pines such as Pinus echinata, but reduced in mature fascicles to 35mm. Leaves in
P. elliottii and P. taeda or with Taxodium distichum. fascicles of 3, usually rigid, or flexible and drooping,
Numerous other angiosperms are components of persisting up to 4 years, straight, (7)913(15)cm
these swamp forests, where waterlogging is only inter- long, 11.2mm wide, serrulate along margins, acute-
700 mittent, most are trees and shrubs or woody vines. pungent, lustrous green. Stomata in several lines on
all faces of the leaves. Pollen cones ovoid-oblong
Conservation to cylindrical, 1.52cm 56mm, yellowish. Seed
cones appearing on very young trees, subterminal or
IUCN: LC lateral, in whorls of 38, sometimes more, on short,
stout, tenacious peduncles, at maturity appearing
Uses sessile, long persistent, serotinous. Mature cones
serotinous, narrowly and obliquely ovoid to ovoid-
The quality of its timber is poor, the wood is brittle attenuate, slightly curved or straight when closed,
and not durable while the scattered occurrence of with an oblique base, (6)813(15) (4)57cm
this species also discourages exploitation on a large when open (width 3.55cm when closed). Seed
scale. Locally it may support a small scale forest scales remaining closed several to many years, part-
industry if mixed with other pines. In some parts of ing eventually or not, oblong, straight or recurved
its range it is planted and harvested for Christmas when parted. Apophyses flat or slightly raised,
trees, but it needs shearing repeatedly to attain the rhombic to obtrullate in outline, with an undulate
desired density of foliage and shape. Its horticultural or crenate upper margin, (weakly) transversely
merits are deemed to be low and it is rarely grown keeled, sometimes gibbous on one side towards base
outside arboreta and botanic gardens. of cone, lustrous ochraceous or light brown, weath-
ering grey. Umbo dorsal, depressed or flat, with a
minute, deciduous prickle. Seeds obliquely obovoid
Pinus greggii Engelm. ex Parl., in Candolle, Prodr. to ellipsoid, slightly flattened, 58 34mm, grey-
16 (2): 396. 1868. brown to blackish brown. Seed wings oblong, with
a straight and a curved side, 1520 5.58mm, yel-
Etymology lowish to grey-brown.
Ecology
Pinus greggii Engelm. ex Parl. var. greggii. Type:
Mexico: Coahuila, San Antonio de las Alanzanas, This variety is generally growing on acid soils. It is
near Saltillo, J. Gregg 402 (lectotype MO). a relatively rare pine that always grows sporadically
among other species of pine, or in mixed pine-oak
Description woodland.
variously serotinous, 35cm wide when closed, fire in the heat of the sun. Although closed stands
47cm wide when opened. Seed scales thick woody, exist, it is more commonly scattered in maquis or
rigid, straight, oblong; apopyses nearly flat or slightly garrigue vegetation on sunny hills and slopes down
raised, weakly transversely keeled and with thin rays to the sea shore, most commonly on limestone
radiating from the centre, more or less rhombic or and dolomite. In stands where fire has been absent
often with a rounded upper margin, to 20mm wide for a longer period, oaks (Quercus suber, Q. ilex)
at mid-cone, lustrous orange-brown or red-brown invade and will eventually dominate. Presumably
weathering grey; umbo flat or depressed, 47mm increased frequency of fire caused by human
wide, broadly rhombic in outline, tan or grey- activities gives the advantage to P. halepensis. Its
brown, unarmed. Seeds obovoid, slightly flattened, altitudinal range is from sea level to ca. 1700m
(5)67(8)mm long, grey-brown, dark mottled; (in Morocco). 703
wing 1525mm long, 812mm wide, more or less
straight, grey-brown with darker streaks. Conservation
Etymology Distribution
This species was named after the plant collector C. T. Mexico: in Chihuahua (Cerro Mohinora), S
Hartweg, who made extensive collections in Mexico Coahuila, S Nuevo Len, Durango, SW Tamaulipas,
in the decade 18381848. Jalisco (Nevado de Colima), Michoacn, Mxico,
Morelos, Hidalgo, Distrito Federal, Tlaxcala, Puebla,
Vernacular names W Veracruz, Guerrero (Cerro Teotepec and vicin-
ity), Oaxaca and Chiapas; in Guatemala in most of
Hartwegs pine; ocote, pino de las alturas (Mexico) the SW highlands; in Honduras on several isolated
mountain summits; reported from the extreme N of
704 Description El Salvador, but not confirmed.
TDWG codes: 79 MXC-DF MXC-ME MXC-MO
Trees to 2530m tall, d.b.h. to 0.01m; trunk mono- MXC-PU MXC-TL MXE-AG MXE-HI MXE-QU MXE-
podial, erect. Bark thick, scaly, deeply fissured, SL MXE-ZA MXG-VC MXN-SI MXS-GR MXS-JA
divided into small or large, square or irregular plates; MXS-MI MXS-NA MXS-OA MXT-CI 80 GUA HON
outer bark dark brown, weathering grey. Branches
spreading and assurging, often persistent, form- Ecology
ing a dense, rounded crown in mature trees. Shoots
thick, rigid, purplish brown, sometimes glaucous. Pinus hartwegii is the typical high altitude pine of
Cataphylls 1520mm long, subulate, recurved, scar- Mexico and Guatemala, where it often forms exten-
ious, brown, weathering blackish grey. Vegetative sive, monotypic pine forests up to the tree line on
buds ovoid-conical; terminal bud up to 30 15mm; high, isolated volcanos or summits of mountain
lateral buds smaller, not resinous. Fascicle sheaths of ranges. In Honduras it is rare, of limited extent and
young leaves 3040mm long, reduced in older fasci- usually found with Abies guatemalensis, Cupressus
cles to (10)1520mm, resinous. Leaves in fascicles lusitanica, Juniperus standleyi, Quercus spp.,
of (3)45(6), most commonly 5, persisting 2(3) Dendropanax lempirianus, Drymis granadensis, a
years, straight or curved, not twisted, rigid, (6)10 ground cover of Ericaceae, Lycopodiaceae, and epi-
17(22)cm long, (1)1.21.5mm wide, with serrulate phytic Bromeliaceae in a cool cloud forest type usu-
margins, acute-pungent, dull light green or glaucous ally between 27002850m on the highest mountain
green. Stomata on all faces of leaves. Pollen ovoid- summits. Similar forests occur in Guatemala and the
oblong to cylindrical, to 2025 67mm, yellow- southern states of Mexico, but there extensive pine
ish or pink-purplish, maturing to brown. Seed cones forests predominate, in which P. hartwegii increas-
subterminal, in pairs or whorls of 36 on short, stout ingly dominates with rising altitude. Its altitudinal
peduncles which are hidden by basal scales. Mature range in Guatemala and Mexico is similar: (2300
cones ovoid-oblong, with acute, curved apex when )25004000(4300)m a.s.l. At lower elevations it
closed, (obliquely) ovoid with a flattened base when is often mixed with P. montezumae, with which it is
opened, 812(14) 58cm. Seed scales spreading closely related, and with other pines depending on
wide, often recurved near base, oblong, thin woody, the geographical area. Soils are both from volcanic
flexible or more rigid. Apophysis more or less flat, and granitic rock, of various depths, but often poor
(weakly) transversely keeled, sometimes gibbous, in nutrients. Climatically there are considerable dif-
rhombic in outline with an angular or irregular upper ferences congruent with latitude/altitude gradients,
margin, varying from light brown to purplish brown with heavy frost and snow during several months
with a blackish centre. Umbo dorsal, depressed, flat and often high winds near the tree lines of the high
or raised, obtuse or with a deciduous prickle. Seeds volcanos in Central Mexico.
broadly ovoid, slightly flattened, 56mm long, light
or dark brown with black spots. Seed wings oblong Conservation
with a straight and a curved side, 1220 712mm,
brown tinged with dark stripes. IUCN: LC
Distribution Pinus henryi Mast., J. Linn. Soc., Bot. 26: 550. 1902.
Pinus massoniana Lamb. var. henryi (Mast.)
Balkan Peninsula, scattered from Bosnia C. L. Wu, Acta Phytotax. Sin. 5 (3): 153. 1956;
Herzegovina to N Greece, Bulgaria (Rhodope Mts.) Pinus tabuliformis Carrire var. henryi (Mast.)
and S Italy (Calabria and Basilicata). C. T. Kuan, Fl. Sichuan. 2: 113. 1983; Pinus tabuli-
TDWG codes: 13 ALB BUL GRC ITA-IT YUG-BH formis Carrire subsp. henryi (Mast.) Businsk, Acta
YUG-CR YUG-KO YUG-MA YUG-MN YUG-SE Pruhoniciana 68: 26. 1999. Type: China: Hubei,
[locality unknown], A. Henry 6909 (holotype K).
Ecology
Pinus massoniana Lamb. var. wulingensis C. J. Qi &
706 Pinus heldreichii is a montane to subalpine species, Q. Z. Lin, Bull. Bot. Res. Harbin 8 (3): 143. 1988.
in Albania, Bosnia and Serbia usually growing above
2200m a.s.l. up to 2640m, but elsewhere down to Etymology
1000m. It is most commonly found on steep moun-
tainsides with very thin soil over limestone, most This pine species was named after Augustine Henry
often in pure, scattered stands. It grows very slowly in (18571930), an English dendrologist who was an
this habitat and presumably ancient specimen trees early botanical visitor to China.
are known, e.g. in Calabria. Although occurring in
the Mediterranean region, its altitudinal range sub- Vernacular names
jects it to sometimes quite severe winter frosts.
Henrys pine; ba shan song (Chinese)
Conservation
Description
IUCN: LC
Trees to 25m tall; trunk to 1m d.b.h., monopodial.
Uses Bark scaly, fissured, breaking into large irregular
plates, greyish brown weathering grey, flaking in
Heldreichs pine is not an important timber tree, small or large chips. Branches spreading or curving
although it is planted in some Balkan countries with down, contorted, forming a domed or flat-topped
timber production as an objective. The wood is used crown. Foliage branches stout, becoming rough with
for heavy construction or as round timber, e.g. for pulvini after shedding leaves, glabrous, new shoots
poles, and to build traditional fences. Its main value reddish brown, usually glaucous, turning light
is as an ornamental tree and it is fairly widely cul- brown to grey-brown or grey. Buds oblong, acute;
tivated for this purpose. It forms as a young tree a terminal bud to 20mm long, slightly resinous; cata-
dense, conical crown and has attractive glaucous to phylls appressed, pale brown. Leaves in fascicles of
light grey shoots and dark green needles. In culti- 2, more or less remote, persisting 23 years, spread-
vation it grows relatively fast, given better condi- ing, held in a 1015mm long, persistent basal sheath,
tions than in its natural habitat. Several cultivars are 712cm long, straight or curved, pliant, 0.71mm
known, usually under the synonym P. leucodermis. wide, often slightly twisted, green; margins minutely
Some of these are slow growing dwarf conifers. serrulate; apex acute or acuminate; stomata in fine
In the Balkans, a number of botanical varieties has lines on all surfaces. Pollen cones in clusters at base
been named under P. heldreichii, it is not known if of new shoots, spirally arranged, short cylindrical,
any of these are in cultivation. ca. 2cm long, yellow. Seed cones solitary or in pairs,
short pedunculate, usually persisting a few years but
opening soon, ovoid when closed, near-symmetri-
cal, 2.55cm long and 2.55cm wide when opened.
Seed scales thin woody, rigid, oblong to obovate,
except basal scales spreading wide, often recurving,
brown. Apophyses slighty raised, variously shaped,
transversely keeled or with 4 radial ridges, ripen- when the opportunity arose during the early decades
ing from green to lustrous brown or reddish brown; of the 20th century.
umbo depressed with a minute prickle. Seeds ovoid-
oblong, 3.55 23mm, pale brown, often mottled;
wing 912mm long, 45mm wide, dark brown. Pinus herrerae Martnez, Anales Inst. Biol. Univ.
Nac. Mxico 11: 76. 1940 [herrerai]. Pinus teocote
Taxonomic notes Schiede ex Schltdl. & Cham. var. herrerae
(Martnez) Silba, Phytologia 68: 63. 1990. Type:
This species has been treated as a variety of Pinus Mexico: Jalisco, Tecalitln, Sierra del Halo,
tabuliformis in Flora of China 4 (1999) and is indeed M. Martnez 3427 (lectotype MEXU).
similar in many characters to that species. Its leaves 707
are not wider than 1mm and its seed cones are Etymology
smaller, with scales that have only slightly raised
apophyses. It was also classified as a variety of This species was named after Prof. D. Alfonso
P. massoniana, from which it differs in its much L. Herrera, a Mexican naturalist.
shorter leaves, its uninodal growth of shoots,
smaller, nearly globose-ovoid seed cones, and rela- Vernacular names
tively shorter seed wings.
Herreras pine; Ocote, Pino chino (Spanish)
Distribution
Description
China: Chongqing, Hubei, Hunan, Shaanxi, Sichuan.
TDWG codes: 36 CHC-CQ CHC-HU CHC-SC Trees to 3035m tall, d.b.h. to 0.81m; trunk mono-
CHN-SA CHS-HN podial, straight or sometimes tortuous. Bark thick,
rough, with scaly plates and shallow, longitudinal
Ecology fissures, reddish brown to grey-brown. Branches
long, slender, arching or spreading horizontally,
Pinus henryi is most common at middle elevations the foliage slightly pendulous. Shoots smooth, with
in the mountains of Central China, and occurs from well developed, long decurrent pulvini, orange-
1100m to 2000m a.s.l. It prefers dry, sunny slopes brown. Cataphylls subulate, scarious, recurving,
and hills, where competition from broad-leaved brown. Vegetative buds ovoid-acute to ovoid-oblong
trees is less severe as the woods are more open and or cylindrical; terminal bud 1015mm long; lat-
lower than in more mesic sites. It is also a pioneer eral buds smaller, not resinous. Fascicle sheaths up
in secondary vegetation, and is there commonly to 20mm long, torn by elongating and spreading
mixed with deciduous shrubs and trees, in later leaves, persistent but reduced to 815mm in mature
stages of the succession often giving way to these fascicles. Leaves in fascicles of 3, persisting 3 years,
angiosperms. slender, lax, drooping or spreading, (10)1520cm
long, 0.70.9mm wide, serrulate at margins, acute,
Conservation light green to yellowish green. Stomata on all faces
of the leaves. Pollen cones spreading, ovoid-oblong
IUCN: NT to cylindrical, 1.51.8cm 5mm, yellowish green,
tinged red. Seed cones subterminal, solitary or
Uses opposite, rarely in whorls of 3, on 1015mm long
peduncles, at first erect, soon on recurved peduncles
This species is similar in its wood properties to P. which fall with cones. Mature cones narrowly ovoid
tabuliformis, but it is less common and widespread, when closed, nearly symmetrical or slightly curved,
and as a consequence of this less important as a tim- ovoid when opened, (2)33.5(4) 23.5cm when
ber tree. Pinus henryi is apperently absent or very open. Seed scales thick woody, oblong, straight or
rare in cultivation; it was not introduced to Europe recurved. Apophysis slightly raised, mostly so on a
few proximal scales on one side of cone, transversely Pinus hwangshanensis W. Y. Hsia, Contr. Inst. Bot.
keeled; apical margin entire or crenate, radially stri- Natl. Acad. Peiping 4: 155. 1936. Pinus luchuensis
ate, light brown. Umbo dorsal, pyramidal, with a Mayr var. hwangshanensis (W. Y. Hsia) C. L. Wu,
minute, deciduous prickle. Seeds (ob)ovoid, slightly Acta Phytotax. Sin. 5 (3): 158. 1956; Pinus luchuen-
flattened, 2.54 23mm, dark grey-brown. Seed sis Mayr subsp. hwangshanensis (W. Y. Hsia) D. Z.
wings obliquely ovate, 58 35mm, translucent, Li, Edinburgh J. Bot. 54 (3): 341. 1997. Type: China:
yellowish with a dark tinge. Anhui, Huang Shan, Si Hai Men, P. C. Tsoong 3111
(holotype PE).
Distribution
Pinus taiwanensis Hayata var. damingshanensis
708 Mexico: along the Sierra Madre Occidental and W. C. Cheng & L. K. Fu, Acta Phytotax. Sin. 13 (4):
more abundantly along the Sierra Madre del Sur; in 85. 1975.
SW Chihuahua, Sinaloa, Durango, W and S Jalisco,
Michoacn and Guerrero. Etymology
TDWG codes: 79 MXE-CU MXE-DU MXN-SI MXS-
GR MXS-JA MXS-MI The species epithet hwangshanensis refers to the
Huang Shan mountain range in S Anhui, China.
Ecology
Vernacular names
The altitudinal range of P. herrerae is (1100)1500
2600 m a.s.l.; its lowest altitude apparently is Huang Shan pine; huang shan song (Chinese)
reached in the Sierra de Cuale in W Jalisco. It occurs
in the mesic forest belt, with annual precipitation Description
of ca. 9001600mm, but with a dry season lasting
from November to May. From N to S annual pre- Trees to 45m tall; trunk to 1m d.b.h, bole straight or
cipitation increases. Mixed pine and pine-oak for- more or less tortuous. Bark on trunk rough and scaly,
est is its usual habitat, in which it is associated with breaking into large plates and deep fissures, grey-
many other pines according to their ranges, further brown to dark grey. First order branches long and
with Pseudotsuga locally. Other broad-leaved trees, spreading, slowly and sometimes only partly self-
e.g. Arbutus, Alnus, Clethra, Juglans, Persea, Clusia, pruning, leaving stumps on the trunk; higher order
and Tilia, are sometimes common. In many areas branches assurgent and densely crowded, forming
the forests are greatly influenced by man-made flat-topped or domed crowns in natural habitat.
changes in their composition, e.g. by selective fell- Foliage branches glabrous, more or less smooth,
ing of larger trees of Pinus, or by recurrent fires light yellowish brown. Buds ovoid-conical to cylin-
and grazing. drical, 1015mm long, 57mm wide, resinous; cata-
phylls appressed, reddish or chestnut brown. Leaves
Conservation in fascicles of 2, held by a persistent, slender basal
sheath 510mm long, straight or slightly curved,
IUCN: LC (5)1017cm long, slender, pliant, slightly twisted,
0.61mm wide; margins minutely serrulate; apex
Uses acute; stomata in fine lines on all surfaces. Pollen
cones clustered, spirally arranged, short cylindri-
Along with other tall growing pines, this species is cal, 1.52cm long, yellow tinged with red, becom-
(heavily) exploited for timber throughout almost ing reddish brown. Seed cones solitary or sometimes
all of its range. It is considered to be of good wood in pairs, persistent on short peduncles, spreading,
quality. Provenance sampling has been started and 36cm long, narrowly ovoid when closed, widen-
is recommended by foresters (Dvorak & Donahue, ing to 2.55cm when open. Seed scales thin woody,
1992) to be conducted on a larger scale. This pine is rigid, oblong, ca, 2.5 1.3cm at mid cone (in larger
unknown in horticulture. cones), straight when spreading, chocolate brown.
Apophysis rhombic in outline or with rounded mon angiosperm (broad-leaved) trees belong to the
upper margin, nearly flat, transversely keeled, Fagaceae. In undisturbed forest the pines would
slightly rugose, lustrous light brown; umbo broadly occupy areas with shallow, sandy acidic soils and
ellipsoid, depressed, armed with a small, persistent rocky outcrops, but in many areas forest disturbance
prickle. Seeds ellipsoid-ovoid, 56mm long, slightly has probably favoured the light demanding pines
flattened; wing 1520mm long, persistent. over broad-leaved trees.
Description Ecology
Trees to 2530(35)m tall, d.b.h. to 6080(100)cm; This species occurs at altitudes between 8001200(
trunk monopodial, straight. Bark moderately thick, 1650)m a.s.l. in mountainous terrain on deep, acidic
rough, scaly with irregular, longitudinal plates and soils derived from granitic rocks. The climate is
shallow fissures, reddish to greyish brown. Branches subtropical, with a 56month dry season from
spreading or ascending; branches of higher orders December to May and annual precipitation from
slender, flexible, slightly pendulous, forming a 10001500mm depending on altitude and exposi-
rounded, rather open crown. Shoots smooth, tion. Pinus jaliscana is a local constituent of pine or
orange-brown, later grey-brown. Cataphylls small, pine-oak forests with a limited extent in most locali-
710 subulate, recurving at apex. Vegetative buds ovoid- ties known so far. More common and widespread
oblong to conical; terminal bud 1015mm long; lat- species that grow with it are P. maximinoi and P.
eral buds ovoid-acute, smaller, not resinous. Fascicle oocarpa, and at slightly higher altitudes also P. doug-
sheaths up to 15mm long, light brown, persistent but lasiana. Several species of Quercus are often codomi-
reduced to 810mm on mature fascicles, weather- nant, Clusia salvinii is also common.
ing light grey. Leaves in fascicles of (4)5, rarely 3,
persisting 23 years, slender, lax, erect or slightly Conservation
drooping, 1218(22)cm long, (0.5)0.60.8mm
wide, with serrulate margins, acute, light green to IUCN: NT
yellowish green. Stomata on all faces of leaves, some-
times inconspicuous. Pollen cones ovoid-oblong to Uses
cylindrical, 1.21.8cm 56mm, purplish yellow,
turning light brown. Seed cones subterminal, soli- No particular preference to single this species out
tary or in whorls of 23 on stout, curved, 715mm as a source of timber is known to occur; it is locally
long peduncles remaining attached to fallen cones. exploited with other pines that grow with it and are
Mature cones ovoid-oblong to ovoid-attenuate when more abundant. This species is not known to be in
closed, often oblique at base, (4.5)68.5(9.8)cm cultivation.
long, (3)45(6)cm wide when open. Seed scales
oblong, straight or slightly reflexed, thick woody.
Apophyses slightly raised to gibbous, more pro- Pinus jeffreyi Balf., in Murray, Bot. Exped. Oregon
nounced on one side of cone towards base, trans- 8: 2. 1853. Type: USA: California, Shasta Co., Shasta
versely keeled, rhombic to pentagonal in outline, Valley, (Chastey Valley, Lat. 41 30 N), J. Jeffrey
with sometimes crenate upper margins, radially 731 (holotype E) & USA: California, Trinity Co.,
striate, lustrous ochraceous or light brown. Umbo west side of Scott Mountain, on Trinity Siskiyou
dorsal, flat to slightly raised, with a minute, decidu- County line, R. O. Alava & A. Alava 2439A
ous prickle. Seeds obovoid, slightly flattened, 3.56 (epitype E).
23.5mm, dark grey-brown. Seed wings obliquely
ovate-oblong, 1317 68mm, yellowish, translu- Pinus jeffreyi Balf. var. baja-californica Silba,
cent, with a grey or black tinge. Phytologia 68: 52. 1990.
Distribution Etymology
Mexico: Jalisco, in the NW part of the Sierra Madre This species was named after John Jeffrey, a plant
del Sur, on the Pacific slope, mainly in the Sierra de collector who discovered it in 1852 in the Shasta
Cuale (Sierra el Tuito) and S of Villa Purificacin. Valley of California.
TDWG codes: 79 MXS-JA
Vernacular names
Description Distribution
Trees to 50m tall, d.b.h. to 1.5m; trunk monopo- USA: California, W Nevada, S Oregon; Mexico: Baja
dial, straight. Bark thick, rough and scaly, divided by California Norte.
deep fissures into thick, elongated plates, light buff TDWG codes: 73 ORE 76 CAL NEV 79 MXN-BC
to light brown or red-brown, the fissures dark grey-
brown. Branches spreading horizontally or assur- Ecology
gent at ends, persistent, forming an open pyramidal
or flat-topped crown. Shoots stout, very rough and Pinus jeffreyi is a montane to subalpine species
scaly, light orange-brown, often glaucous. Cataphylls largely confined to the mountains of California, with
ca. 20 mm long, triangular-subulate, reflexed, an altitudinal range of (50)300m to 3050m a.s.l. 711
scarious, with erose-ciliate margins, light brown. It is tolerant of low temperatures in winter and can
Vegetative buds ovoid-acute or subglobose; terminal grow on thin soil or even in crevices of bare granite
bud 1520mm long; lateral buds smaller, not resin- rock. In the Sierra Nevada of California the species,
ous. Fascicle sheaths to 2025mm long, persistent with its close relative P. ponderosa, is characteris-
but reduced to 1015mm in older fascicles. Leaves in tic of open, dry and summer-warm mixed forests
fascicles of 3, sometimes a few fascicles of 2, persist- of the Yellow pine belt where it tends to occupy
ing (3)45(6) years, thick, rigid, straight or slightly the upper zone towards the tree line. In the south-
curved, sometimes twisted, (12)1522(25) cm ern part of this mountain range it occurs in diverse
long, 1.51.9(2)mm wide, with serrulate margins, mixed coniferous forest with e.g. Pinus ponderosa,
acute-pungent to acuminate, light yellowish green to P. lambertiana, P. monticola, P. contorta, Abies con-
greyish green. Stomata on all faces of leaves, in con- color, A. magnifica, Calocedrus decurrens, Juniperus
spicuous lines. Pollen cones ovoid-oblong to cylin- occidentalis, and Sequoiadendron giganteum. In
drical, 23.5cm 67mm, yellow or purplish yellow. southern California and Baja California only Abies
Seed cones subterminal, solitary or in pairs on short, concolor, Calocedrus decurrens, Pinus contorta, and
stout, persistent peduncles, spreading and seemingly P. lambertiana accompany P. jeffreyi. In the Klamath
sessile at maturity, leaving a few basal scales on the Mountains of Oregon P. jeffreyi occurs on thin ultra-
branch when falling. Mature cones broadly ovoid to mafic soils of volcanic origin (peridotites and ser-
subglobose, with a slightly oblique, flattened base, pentine) which are poor in nutrients; on these soils
(10)1225(30) 914(16)cm when open. Seed its most common associate is Calocedrus decurrens.
scales parting soon and wide, thin woody, straight Here it descends to low elevations (around 100m),
or slightly curved, up to 20mm wide. Apophysis while in the Sierra Nevada it ascends to 2900m and
slightly raised, transversely keeled, broadly rhom- in the Sierra San Pedro Martr of Baja California to
bic in outline, on the proximal scales more gibbous, 3050m.
often resinous, ochraceous to light brown. Umbo
dorsal, moderately raised, transversely keeled, ter- Conservation
minating in a distinct, curved, 35mm long spine.
Seeds obliquely ovoid, slightly flattened, 912mm IUCN: LC
long, light yellowish brown with faintly darker spots.
Seed wings oblique, with a straight and a curved Uses
side, 2025 10mm, light yellowish brown to light
brown. Jeffrey pine is very similar to Ponderosa pine in its
wood properties and is consequently an important
Taxonomic notes timber tree. It has a preference for poorer sites and
higher altitudes, and consequently grows slower than
Pinus jeffreyi is closely related to P. ponderosa and P. Ponderosa pine in its natural habitat, but it grows
coulteri and introgression via pollen may occasion- as fast or faster in plantations. The timber industry
ally occur where these species occur together. does not differentiate the wood from the two spe-
cies; details are therefore given under P. ponderosa.
The resin of P. jeffreyi contains much heptane and 0.7(1)mm wide; margins minutely serrulate; apex
lower levels of terpenes than found in other pines; in long acuminate; stomata in fine lines on all surfaces.
the past this biochemical product was distilled and Pollen cones clustered and spirally arranged at base
used as an additive to raise the octane levels of pet- of new shoots, short cylindrical, yellow. Seed cones
rol (gasoline), as well as for medicinal applications. solitary or in pairs or threes, short pedunculate,
Jeffrey pine is not uncommon in cultivation, but it spreading or reflexed, persistent, 4.57(8)cm long,
is mostly restricted to large parks and gardens with narrowly ovoid-conical when closed, symmetrical
tree collections (arboreta). In forestry, crosses and or slightly non-symmetrical, ovoid to subglobose
back-crosses have been experimented with a num- when opened, then 34(5.5)cm wide. Seed scales
ber of related pines, in particular P. coulteri and P. woody, rigid, more or less recurved when spread-
712 ponderosa and its varieties. ing, narrowly oblong, 2.53 11.5cm, dark brown.
Apophysis prominently raised, more or less pyra-
midal, irregularly rhombic in outline, transversely
Pinus kesiya Royle ex Gordon, Gard. Mag. & Reg. keeled, light brown. Umbo small, ellipsoid or at
Rural Domest. Improv. 16: 8. 1840. least protruding, armed with a very small, persis-
tent prickle. Seeds ellipsoid, 57(8)mm long,
Etymology slightly appressed, dark brown to nearly black; wing
1520mm long, 68mm wide, persistent.
The species epithet refers to the Khasi Hills in NE
India, from where it was first reported by J. F. Royle; Distribution
its spelling has long been a matter of unnecessesary
controversy. Royles (European) spelling of the top- China: Yunnan; NE India: Assam; Indochina;
onym at the time was as good as any other and must Malesia: Philippines (Luzon).
be followed for the name of this pine in botanical TDWG codes: 36 CHC-YN 40 ASS-AS ASS-MA
nomenclature. ASS-ME ASS-MI ASS-NA 41 CBD LAO MYA THA VIE
42 PHI
Vernacular names
Ecology
Khasia pine, Khasi pine, Luzon pine, Benguet pine;
dingsa, far, saral (India); tinyu (Myanmar). Pinus kesiya occurs in pine savannas, pure stands
with nearly closed canopy, and mixed pine-broad-
Description leaved forests in valleys with e.g. Quercus serrata
and Alnus nepalensis along streams. It occupies
Trees to 3035(45)m tall; trunk to 1m d.b.h., often drier sites in NE India, Myanmar and Thailand, at
straight and columnar. Bark on trunk thick, deeply altitudes generally between 800 and 1500m a.s.l.,
fissured forming irregular plates, rough and flaking, occasionally to 2000m. Further east, in Lao PDR,
brown weathering grey-brown. Branches spread- Viet Nam and on the island of Luzon in the
ing, usually crooked, slowly self-pruning, forming Philippines, its altitudinal range is greater and it is
a broadly domed crown. Foliage branches slender, found up to 27003000m a.s.l. in a much wetter
rough with pulvini from fallen leaf fascicles, later climate. Despite this, fires are frequent, creating an
more or less smooth as these disappear, lustrous yel- open, grass-dominated woodland or savanna with
lowish brown, darkening to orange-brown, shoots scattered stands of pines or solitary trees. The soils
multi-nodal within one year. Buds small, conical, are usually sandy or loamy and derived from sand-
without resin or slightly resinous; cataphylls red- stone or quarzite. Pinus kesiya is a pioneer in defor-
brown. Leaves in fascicles of 3, held together in a ested secondary vegetation, especially if fire has been
1520mm long, persistent basal sheath, (10)12 a factor in the disturbance.
22(25)cm long, slender, very lax or flexible, 0.5
Etymology Distribution
The species epithet refers to Korea. China: Heilongjiang, Jilin, Liaoning; Japan:
Hokkaido, Honshu; North & South Korea; Russian
Vernacular names Far East: Amur, Khabarovsk, Primorye.
TDWG codes: 31 AMU KHA PRM 36 CHM-HJ CHM-JL
Korean pine; chas namu (Korea, DPR); hong song CHM-LN 38 JAP-HK JAP-HN KOR-NK KOR-SK
(Chinese)
Ecology
Description
714 In the Russian Far East this pine grows from 200m
Trees to 2530m tall (in China reported to attain to 600m a.s.l., 500m to 1300m a.s.l. in China, and in
50m); trunk to 1m d.b.h., growing to a straight, Japan it occurs to an altitude of 2500m. The climate
columnar bole. Bark smooth in young trees, in has a summer monsoon character within proxim-
larger trees becoming rough and scaly, with shal- ity of the coast, but with a strong continental influ-
low grooves and longitudinal fissures, forming ence further inland. Temperature extremes range
oblong, reddish grey or grey-brown plates; inner from +37 C to 45 C within its natural range. Pinus
bark red-brown. Branches spreading or ascending, koraiensis grows in dry places on podzols among
forming a broad crown; top of trees often forked. deciduous broad-leaved trees like oaks, poplars and
Foliage branchlets slender, new shoots occasion- birches, but on the Russian coast of the Sea of Japan
ally pubescent with yellowish hairs, red-brown it is codominant with Abies holophylla, forming
becoming grey-brown. Buds oblong or ovoid, the groves of conifers in a more varied deciduous broad-
central bud to 10mm long, slightly resinous; cata- leaved forest. In Japan it also occurs together with
phylls reddish brown. Leaves in fascicles of 5, per- other pines. In Korea and NE China (Manchuria)
sisting 23 years, held in a deciduous basal sheath this pine has been heavily exploited, resulting in the
of orange-brown flimsy scales, slender, straight or disappearance of many magnificent pine forests.
abruptly bent, 613cm long, pliant or lax, ca. 1mm
wide, green on abaxial surface; margins minutely Conservation
serrulate; stomata in bluish white lines on the two
adaxial faces. Pollen cones in small clusters at base Despite over-exploitation resulting in removal of
of new shoots, short ellipsoid-cylindrical, ca. 15mm many indigenous forests with this species, its wide
long, yellow. Seed cones solitary or in whorls of 24 distribution and partial regeneration (outside affor-
on stout, 11.5cm long peduncles, semi-persistent, estation) appears to guarantee the survival of this
initially erect, becoming spreading to pendulous as species, which is therefore not considered threat-
they grow, blue-green and resinous, when mature ened under IUCN criteria.
914cm long and 68cm wide, ovoid-oblong, light IUCN: LC
brown or grey-brown. Seed scales at base of cones
reflexed or recurved, more or less indehiscent to Uses
slightly spreading but not releasing the seeds, more
or less rhombic in outline, the middle scales ca. 3cm Korean pine is a highly important timber tree; in
long, 2.5cm wide at transition to apophysis, with large parts of its range it has been over-exploited,
two deep seed cavities adaxially; apophyses broadly but it is now used widely in afforestation schemes
triangular, slightly thickened, oblong at cone apex, especially in NE China. Its timber is of good quality,
longitudinally striated or grooved, nearly straight light and soft, straight grained and easy to work with
but recurved or reflexed at the margin towards in milling and carpentry. It is fairly decay resistant
apex; umbo terminal, obtuse or acute. Seeds tri- and therefore finds uses like telephone poles, rail-
angular-obovoid, 1216mm long, light brown or way sleepers, wooden bridges, and boat building. In
grey-brown, without a wing when removed from construction it provides building timbers as well as
the scale. flooring, plywood and veneers. It can be chipped for
715
Plate 30. Pinus krempfii. 1. Habit of tree. 2. Branch with foliage. 3. Leaves. 4. Seed cone. 5. Seeds.
particleboard or flakeboard manufacture, or pulped few thin, acute scales which fall separately, spreading,
for the paper industry. More specialized uses are fur- straight or slightly curved, 45(7) cm long,
niture, sports equipment and musical instruments. extremely flat, 1.55mm wide, often twisted in the
Resin is extracted from wood pulp and used to pro- lower, narrower fourth to third of their length, with
duce turpentine and other products. The seeds are a narrow median rib on each side, gradually taper-
rich in vegetative oil with a high nutritive value, and ing to an acute apex, green on adaxial side, glau-
this is used in the food processing industry; seeds can cous green on abaxial side; stomata in fine lines on
also be consumed whole and most of the imported abaxial face. Pollen cones small, in groups at base
pine kernels in Europe and the USA are now sourced of new shoots, 7mm long when full-grown. Seed
from this species. It is widely cultivated as an orna- cones solitary or more commonly in pairs on short
716 mental tree in China, Korea and Japan, but less com- peduncles, maturing within 2 years, spreading and
mon in Europe and the US, where until recently only opening, falling after seed dispersal, 57cm long,
a limited number of cultivars was known, but now narrowly ovoid when closed, light green, opening to
on the increase. 35cm wide, becoming brown. Seed scales relatively
few, soft woody, oblong-concave, dark brown when
mature; apophyses hard woody, thickened, raised
Pinus krempfii Lecomte, Bull. Mus. Hist. Nat. and often curved, prominently transversely keeled,
(Paris) 27: 191. 1921. Ducampopinus krempfii more or less rhombic in outline or with rounded
(Lecomte) A. Chev., Rev. Int. Bot. Appl. Agric. upper margins, light brown or light reddish brown;
Trop. 24: 30. 1944. Type: Viet Nam: South umbo dorsal, raised, acute or more or less flat with
Viet Nam, Tinh Khanh Hoa, near Nha Trang, a minute prickle. Seeds very small, 2.53mm long,
M. Krempf 1537 (holotype P). Pl. 30 more or less ovoid, nearly black; wing long and nar-
row, 1720mm long, 45mm wide, detachable from
Pinus krempfii Lecomte var. poilanei Lecomte, Bull. the seed.
Mus. Hist. Nat. (Paris) 30: 325. 1924.
Taxonomic notes
Etymology
This remarkable pine has long been considered dis-
This species was named after the collector of the type tinct from all other species, even to the extent that it
specimen, M. Krempf. should be classified in its own genus (or subgenus)
Ducampopinus A. Chev. This opinion was mostly
Vernacular names based on the peculiar, flat needles; virtually all other
organs, including the dwarf shoots that produce
Krempf s pine; Thng l dt (Vietnamese) the needles, are matched by other pines and are
mostly unique to the genus. More recent phyloge-
Description netic analyses based on DNA data have given results
that do not support this major distinction and have
Trees to 40(50)m tall; trunk to 3(4)m d.b.h., erect, placed the species firmly within the genus Pinus and
straight, massive, often butressed. Bark smooth on as almost certainly belonging to subgenus Strobus
young trees, becoming rough and scaly on large (Liston et al. in Mill, 2003). It is definitely an ancient
trunks, with irregular plates divided by shallow fis- relict species, as are several other species in this
sures, grey-brown to grey. Branches long, spreading subgenus.
wide and near the top ascending, forming a broadly
domed or flat-topped, dense or more open crown. Distribution
Foliage branches slender, multinodal, smooth after
leaves have fallen, light brown turning grey with age. S Viet Nam (southern Truong Son Range, between
Buds small, almost continuously producing new Dalat and Nhatrang and N of Nhatrang).
shoots and leaves, not resinous. Leaves in fascicles of TDWG codes: 41 VIE
2, with early deciduous basal sheaths consisting of a
718
Plate 31. Pinus lambertiana. 1. Habit of tree. 2. Branchlet with leaves. 3. Leaves. 4. Seed cone.
often more reflexed, angular-obovate to obtrullate, resticted to the highest mountain summits further
up to 40mm wide and 45mm thick. Apophysis south and the diversity of conifers dimishes as does
triangular to obtusely rhombic, 58mm thick at their size.
base, straight, smooth or radially striated, yellowish
brown, often with yellow resin blobs. Umbo termi- Conservation
nal, obtuse-triangular. Seeds obovoid or obliquely
obovoid, (10)1215(18)mm long, 610mm wide, IUCN: LC
dark brown. Seed wings broadly obovoid to dolabri-
form or truncate, 2030mm long, 1215mm wide, Uses
light brown.
Sugar pine is the most valuable of the pines due to 719
Distribution its enormous size and its light, soft, even-grained,
knot-free wood of which very large, straight pieces
USA: California, Oregon, Nevada (extreme western can be sawn. Consequently, old growth stands of
part); NW Mexico: Baja California Norte (Sierra San this species command very high prizes; fortunately
Pedro Martr). many are now protected in National Parks and other
TDWG codes: 73 ORE 76 CAL NEV 79 MXN-BC reserves. This species is used for high quality con-
struction timber and the finished milled wood of
Ecology this king of pines as John Muir called it, makes it
ideal for high standard windows and doors as well
This majestic pine is common in the mixed and var- as foundry casting and even musical instruments
ied conifer forests of the Transition Zone in the high such as organ pipes and piano keys. Plantation for-
mountains of Oregon and California commonly estry has not been very successful and in Europe few
between 600m and 2400m a.s.l., but reaching trees have survived for long in arboreta and parks
28003200m in the south of its range. In the Sierra due to white pine blister rust (Cronartium ribicola,
Nevada of California it is restricted to the western Basidiomycota) to which it is exceptionally sensis-
slopes at middle elevation, between the warmer tive. The vernacular name refers to the sugar content
Upper Sonoran Zone and the colder Canadian Zone. of the resin; in the past native tribes used it as a chew-
Here one of the most impressive conifer forests in the ing gum. This use was first noted by David Douglas,
world occurs, dominated by gigantic trees; indeed who thought the native people were eating it.
the largest tree species in the world, Sequoiadendron
giganteum, is restricted to this zone and this moun-
tain range. Other species are Pinus monticola, Pinus latteri Mason, J. Asiat. Soc. Bengal. Sci. 18:
P.ponderosa, P. jeffreyi, P. contorta var. murrayana, 74. 1849. Pinus merkusii Jungh. & de Vriese var.
Abies magnifica, A. procera, Calocedrus decurrens, latteri (Mason) Silba, Phytologia 68: 53. 1990; Pinus
and in the north of the range Pseudotsuga menzie- merkusii Jungh. & de Vriese subsp. latteri (Mason)
sii. A hospitable climate with warm, sunny sum- D. Z. Li, Edinburgh J. Bot. 54 (3): 346. 1997. Type
mers moistened by rain showers and snowy but not not designated. Fig. 227
extremely cold winters marks this zone. In Oregon
Pseudotsuga menziesii, Abies grandis, Tsuga hetero- Etymology
phylla, Thuja plicata, and Calocedrus decurrens are
its most common associate conifers. Broad-leaved This species was named by the Rev. F. Mason after
trees are rare and much smaller, Arbutus menziesii Capt. Latter, the discoverer.
and Quercus kelloggii are perhaps the most com-
mon. The ground is open, littered with conifer nee- Vernacular names
dles, interspersed with small, flowery meadows and
lush streamsides full of flowering shrubs. The bright Tenasserim pine; shaja, tinshu (Burmese); son-song-
yellow lichen Letharia vulpina festoons trunks and bai (Thai); nan ya song (Chinese)
branches everywhere. This type of forest becomes
charcoal burning is a more traditional use still cur- straight, rigid, 1220(25)cm long, 1.01.2(1.5)mm
rent. It is also an important tree for resin tapping wide, serrulate at margins, acute-pungent, glaucous
in some countries. In China, the resin of this spe- green. Stomata on all faces of the leaves. Pollen cones
cies has some medicinal applications, e.g. in making ovoid-oblong to cylindrical, 12cm 56mm, yel-
ointments. This tropical pine has been introduced lowish green, turning light brown. Seed cones sub-
for afforestation in several countries in Africa, but terminal, sometimes solitary, commonly opposite
the results have on the whole been unsatisfactory on short, stout, curved peduncles which usually
because of difficulties with raising it beyond the fall with cone. Mature cones narrowly ovoid to
seedling stage in all situations where competition ovoid-attenuate when closed, more or less asym-
from other plants (weeds) is a factor. metrical, ovoid when open, with a more or less flat-
tened base, then 58(9) 46(7)cm. Seed scales 721
thick woody, oblong, recurved when fully spread.
Pinus lawsonii Roezl ex Gordon, Pinetum Suppl.: Apophysis slightly raised, transversely keeled, rhom-
64. 1862 [lawsoni]. Type: Mexico: Michoacan, bic in outline, with undulate-crenate upper margin,
Uruapn, Cerro de la Cruz, C. G. Pringle 10141 radially striate, light brown to grey-brown. Umbo
(neotype NY). dorsal, pyramidal, curved, lacking a distinct prickle.
Seeds obovoid, slightly flattened, 45mm long, dark
Pinus lawsonii Roezl ex Gordon var. gracilis Debreczy brown. Seed wings 1216 56mm, translucent,
& Rcz, Phytologia 78 (3): 19. 1995. light brown with a dark tinge.
Etymology Distribution
Roezl named this species after the English gardener Mexico: Michoacn, Mxico, Morelos, Distrito
and plantsman Charles Lawson (17941873), who Federal, Guanajuato, (one locality in) Veracruz,
validated David Douglas name Pinus ponderosa in Guerrero and Oaxaca.
the same way as did George Gordon with this name TDWG codes: 79 MXC-DF MXC-ME MXC-MO
from him. MXE-GU MXG-VC MXS-GR MXS-MI MXS-OA
Lawsons pine; pino ortiguillo (Spanish) Pinus lawsonii has an altitudinal range of 1300
2600m a.s.l. It is a constituent of warm-temperate to
Description temperate montane forest or woodland, with annual
precipitation ranging from 6001500mm and a dry
Trees to 2530m, d.b.h. to 75cm; trunk monopo- season from November to May. It is most commonly
dial, straight, sometimes tortuous. Bark thick, rough found in pine-oak forest, also in pine forest with e.g.
and scaly, with broad and deep longitudinal fissures; Pinus pringlei, P. patula, P. montezumae, P. oocarpa,
outer bark dark blackish brown. Branches spread- P. leiophylla, P. herrerae, P. teocote, and P. pseudostrobus.
ing horizontally, assurgent in the higher part of the On sites with sandy, shallow soil Juniperus can be
crown; forming a broadly domed but open and irreg- codominant.
ular crown. Shoots smooth, orange-brown, often
glaucous. Cataphylls subulate-lanceolate, ca. 10mm Conservation
long, scarious, erose-ciliate at margins, dull brown.
Vegetative buds ovoid-oblong to cylindrical; termi- IUCN: LC
nal bud 1015mm long; lateral buds shorter, acute,
not resinous. Fascicle sheaths of young leaves up to Uses
25mm long, persistent but reduced to 1015mm in
older fascicles. Leaves in fascicles of 34(5), very Along with other pines, which are often more abun-
seldom 2, persisting 23 years, straight or nearly dant in the mixed pine or pine-oak forests, this spe-
cies is cut commercially for lumber, although most Mature cones (narrowly) ovoid when closed, nearly
trees are of medium size and the boles sometimes symmetrical, (broadly) ovoid when opened, (4)5
tortuous. In several places with abundant trees 7(8) (3)45.5cm when open. Seed scales straight
of this species, resin may be tapped as well. As an or recurved near base of cone, oblong, with nearly
ornamental tree this species will only grow success- straight margins. Apophysis raised, transversely
fully in countries with very mild climate; it is not keeled, with the central section of the second sea-
known to be in cultivation for this purpose. sons growth distinctly marked as a narrow band
around the umbo, rhombic in outline, dull brown to
grey-brown. Umbo dorsal, pyramidal, with a blunt
Pinus leiophylla Schiede ex Schltdl. & Cham., prickle, darker than the apophysis. Seeds obliquely
722 Linnaea 6: 354. 1831. ovoid, slightly flattened, 34(5)mm long, dark
grey-brown with black spots. Seed wings obliquely
Etymology oval, 1018 48mm, yellowish brown, translucent,
with a dark tinge.
The species epithet is derived from Latin: lenis = soft
and Greek: phyllos = leaf. Distribution
Along with many other species of pine in the Sierra Pinus leiophylla Schiede ex Schltdl. & Cham. var.
Madre Occidental, Pinus leiophylla has been heavily chihuahuana (Engelm.) Shaw, [Pines Mexico] Publ.
exploited for timber in the latter half of the 20th cen- Arnold Arbor. 1: 14. 1909. Pinus chihuahuana
tury. Due to its high resin content it does not provide Engelm., in Wislizenus, Mem. Tour N. Mexico:
high quality wood but at the same time is a producer 103. 1848; Pinus leiophylla Schiede ex Schltdl. &
of good resin, for which it is tapped. The wood is used Cham. subsp. chihuahuana (Engelm.) E. Murray,
in heavy construction and for crates and boxes. The Kalmia 12: 23. 1982. Type: Mexico: Chihuahua,
variety chihuahuana is less exploited for timber, but Cosiquiriachi, F. A. Wislizenus 232 (holotype MO).
an equally prolific producer of resin. Smooth-leaved
pine has been planted in various parts of the world, Description
especially in Africa, as a plantation forestry tree.
Trees to 1525m tall, often not more than 10m, d.b.h.
2 varieties are recognized: to 2060cm. Shoots reddish brown, often glaucous.
Leaves in fascicles of (2)3(4, rarely 5), (4)612
(14)cm long, 0.91.3(1.5)mm wide. Stomata in
Pinus leiophylla Schiede ex Schltdl. & Cham. var. (4)58(9) lines on the convex abaxial face, in 34
leiophylla. Type: Mexico: Veracruz, Cruz Blanca, lines on each adaxial face. Resin ducts in the leaves
[inter Cruz Blanca et Jalacingo], C. J. W. Schiede (3)46(7), medial, occasionally 12 internal.
1109 (holotype HAL).
Distribution
Pinus lumholtzii B. L. Rob & Fernald var. microphylla
Carvajal, Phytologia 59: 135. 1986. USA: SE Arizona, SW New Mexico; Mexico:
along the Sierra Madre Occidental in NE Sonora,
Description W Chihuahua, Durango, Nayarit, N Jalisco and
Zacatecas.
Trees to 2030(35)m tall, d.b.h. to 5085cm. TDWG codes: 76 ARI 77 NWM 79 MXE-CU MXE-DU
Shoots reddish brown, sometimes glaucous. Leaves MXE-ZA MXN-SO MXS-JA MXS-NA
in fascicles of (4)5(6), 4more often than 6,
(6)815(17)cm long, 0.50.9mm wide. Stomata Conservation
in (3)46(7) lines on the convex abaxial face, in
(2)34 lines on each adaxial face. Resin ducts in the IUCN: LC
leaves (1)23(4), medial, occasionally 1 internal.
Pinus longaeva D. K. Bailey, Ann. Missouri Bot. peltate, the distal part cordate to rounded, smooth,
Gard. 57: 243. 1971. Pinus aristata Engelm. var. lon- ca. 1mm wide. Seed cones subterminal, solitary or
gaeva (D. K. Bailey) Little, Phytologia 42: 221. 1979; in pairs, nearly sessile, falling after seed dispersal.
Pinus balfouriana Balf. subsp. longaeva (D. K. Bailey) Immature cones erect, ovoid, dark purple (some-
E. Murray, Kalmia 12: 23. 1982; Pinus aristata times green), maturing in two seasons. Mature cones
Engelm. subsp. longaeva (D. K. Bailey) E. Murray, ovoid-cylindric becoming more ovoid when open,
Kalmia 13: 21. 1983. Type: USA: Nevada, Wheeler 610 46cm. Apophyses on seed scales thick, tri-
Peak Scenic Area, Wheeler Peak, D. K. Bailey & angular to rhombic, transversely keeled, recurved,
J. E. Whitson 7001 (holotype COLO). Fig. 228 red-brown. Umbo dorsal, transverse-triangular at
base, terminating in a slender but rigid, variable
724 Etymology prickle 16mm long. Seeds ellipsoid or obovoid,
58 35mm, light brown with darker red-brown
The species epithet longaeva refers to the longivity specks. Seed wings 1013mm long.
of this pine.
Distribution
Vernacular names
USA: E California, Nevada, Utah.
Great Basin Bristlecone pine, Intermountain TDWG codes: 76 CAL NEV UTA
Bristlecone pine
Ecology
Description
Like its relatives Pinus aristata and P. balfouriana, P.
Trees, small to medium, height to 16 m; trunk d.b.h. to longaeva is a subalpine to alpine species and like the
3m. Trunk monopodial, straight or contorted, often others a xerophyte, adapted to extreme conditions
strongly tapering in old trees or sometimes multi- and fluctuations of temperature and moisture. Its
stemmed, branching very low and much reduced in altitudinal range is between 2130 and 3700m above
size at tree line. Bark thin, shallowly to deeply fis- sea level and it often grows with P. flexilis, but with
sured, breaking into irregular, scaly ridges, reddish few other trees or shrubs, and often in bare, rocky
brown, weathering grey. Branches of first order con- ground. Weather conditions are often extreme in all
torted, ascending or spreading, persistent, of higher seasons, ranging from heat, extreme radiation and
orders flexible and often pendent. Shoots puberu- desiccating winds to snow and ice storms that blast
lent, with short, non-decurrent pulvini, pale red- the bark off the trunks. Growth is therefore exceed-
brown, ageing to yellowish grey or grey. Cataphylls ingly slow, producing very dense and hard wood
57mm long, subulate, scarious, brown, with entire with annual increments of minimal width, often
margins, soon deciduous. Vegetative buds ovoid and restricted to only parts of the trunk and some of the
acute, resinous; terminal bud 810mm long; lateral branches, the remainder being already dead wood.
buds smaller, all with imbricate, appressed, subu- Yet these trees even when 4000 years old still pro-
late, red-brown scales. Fascicle sheaths deciduous duce cones with viable seeds, and seedlings and sap-
and absent in second years fascicles. Leaves in fas- lings can be found growing around them.
cicles of 5, in dense tufts persisting up to 30 years,
mostly connivent, curved forward against shoot, Conservation
(1.5)23.5(4)cm long, 0.81.2mm wide, lacking
resin droplets or only a few present; margins entire Pinus longaeva was considered vulnerable to extinc-
or distally serrulate; apex acute or short acuminate; tion because of its fragmented occurrence in a limited
leaf colour deep green on the smooth abaxial face, number of populations, often with only a few hun-
white stomatal lines on both adaxial faces. Pollen dred or at most 5001000mature individual trees.
cones crowded at the proximal end of a new shoot, Recruitment is very slow and it seemed uncertain,
spreading, subtended by light brown bracts, ellip- given the great age of these trees, whether this con-
soid or ovoid, ca. 10mm long, red or purple (some- stitutes sufficient replacement of (eventually) dying
times yellow), maturing to brown. Microsporophylls old trees. Recently, a re-assessment by USDA forest
very wind-exposed slopes or close to the sea, where nal bud ca. 15 8mm; lateral buds smaller, resinous.
it becomes wind distorted. This species is very tol- Fascicle sheaths (20)2535mm long on juvenile
erant of saline air and ocean spray, but grows well fascicles, consisting of ca. 10 reddish brown scales,
in planted situations away from strong winds and soon coming apart, forming a tuft of curling scales at
airborne salt. In its natural habitat it mostly forms the base of fascicles before they fall at maturity, leav-
pure, open stands with a low vegetation of grasses ing fascicles bare. Leaves in fascicles of 3 (exception-
and shrubs stabilizing drifting sand, virtually free ally 2 or 4), usually extremely pendulous, persisting
from competition by other trees. two years, lax but not thin, (15)2030(40+)cm
long, (1)1.21.5mm wide, with serrulate margins,
Conservation acute, light green. Stomata on all faces of leaves, in
726 conspicuous lines. Pollen cones sparsely clustered,
IUCN: LC cylindrical, 23cm 5mm when mature, at first
pinkish, turning yellow. Seed cones subterminal or
Uses lateral, solitary (sometimes in whorls of 2, rarely 3)
on 1015mm long, curved peduncles, breaking off
This pine is not important as a timber resource and easily and remaining with the soon deciduous cones.
as such only of local use. It is well adapted to salt- Mature cones ovoid to ovoid-attenuate when closed,
laden sea winds and has been planted to stablize ovoid to ovoid-acute when opened, then (3)3.5
sand dunes in coastal areas in other parts of Japan 5.5(7) (2.5)34.5cm. Seed scales thick woody,
as well as in Taiwan. It is not known to be used as an oblong, spreading wide or lower scales remaining
ornamental tree and no cultivars are known of this closed, with smaller basal scales remaining con-
species. nate. Apophyses slightly raised, thickened along the
distal margin, obscurely transversely keeled, those
on basal scales gibbous, rhombic to pentagonal in
Pinus lumholtzii B. L. Rob & Fernald, Proc. Amer. outline, ochraceous to reddish brown. Umbo dor-
Acad. Arts 30: 122. 1894. Type: Mexico: Chihuahua, sal, often tilted towards distal margin of apophysis,
Coloradas, C. V. Hartman 541 (holotype GH). flat or slightly raised, with a minute, soon deciduous
prickle. Seeds obliquely obovoid, slightly flattened,
Etymology 35mm long, dark brown, often with black dots.
Seed wings oblique, widest below apex, (8)1014
This species was named after the plant collector Carl 46mm, light yellowish or greyish brown.
Lumholtz (fl. 1900).
Distribution
Vernacular names
Mexico: Sierra Madre Occidental, in Chihuahua,
Lumholtzs pine; pino triste (Spanish) Sinaloa, Durango, Nayarit, Jalisco, Zacatecas,
Aguascalientes and Guanajuato.
Description TDWG codes: 79 MXE-AG MXE-CU MXE-DU
MXE-GU MXE-ZA MXN-SI MXS-JA MXS-NA
Trees to 20m tall, d.b.h. to 5070cm; trunk mono-
podial, straight. Bark thick, scaly, very rough, Ecology
divided into elongated, irregular plates and deep,
wide, longitudinal fissures, greyish brown to dark The altitudinal range of this species is (1500)1700
grey. Branches spreading or ascending, branches 2600(2900)m a.s.l., which corresponds to the
of higher orders slender, flexible, drooping. Crown lower and middle slopes of the Sierra Madre. It
broad, rounded and usually open. Shoots ini- grows usually mixed with several species of Quercus
tially glaucous, turning reddish brown, then grey. and other pines in pine-oak forest, or on the wet-
Cataphylls subulate, up to 10mm long, scarious, red- ter western slopes of the Sierra Madre, in mixed
brown. Vegetative buds ovoid-conical, acute; termi- pine forest. Associated pines are e.g. P. leiophylla,
Pinus luzmariae Prez de la Rosa, Bol. Inst. Bot. Similar to that of Pinus oocarpa.
(Guadalajara) 5 (13): 127. 1998. Pinus oocarpa
Schiede ex Schltdl. f. trifoliata Martnez, Anales Conservation
Inst. Biol. Univ. Nac. Mxico 16: 297. 1945; Pinus
oocarpa Schiede ex Schltdl. var. trifoliata Martnez, IUCN: LC
Pinos Mexic., ed. 2: 308, f. 252253. 1948. Type:
Mexico: Durango, Sierra de Chavarria, M. Martnez Uses
3458 (holotype MEXU).
No uses have been recorded, but it will be used in the
Etymology same way as Pinus oocarpa.
No common names have been recorded for this little This species commemorates the botanist Francis
known species. Masson (17411805) who never visited China and is
best known for his plant collecting at the Cape of
Description Good Hope in South Africa in the 1770s.
Pinus massoniana is a pine which grows as well in the 2 varieties are recognized:
lowlands as in the highlands, and occurs from a few
Pinus massoniana Lamb. var. massoniana. Type: may be much overlap of these continuous measure-
Illustration in Lambert, Descr. Pinus 1: t. 12. 1803 ments with cones of var. massoniana on the main-
(lectotype). land. This variety is here only tentatively accepted.
Etymology
Pinus massoniana Lamb. var. hainanensis
W. C. Cheng & L. K. Fu, Acta Phytotax. Sin. 13 (4): The species epithet commemorates the Mexican bot-
85. 1975. Type: China: Hainan, Wang Chin 3117 anist Maximino Martnez (18881964), who studied
(holotype PE). the conifers of Mexico, but did not know this pine; a
concomitant meaning refers to the size (Latin: max-
Description ime) of the cones and seeds.
green, turning orange-brown to grey. Cataphylls ca. soils of igneous origin. Precipitation is probably ca.
5mm long, narrowly triangular-caudate, recurved. 700800mm annually, virtually restricted to four
Vegetative buds small, ovoid-conical; terminal buds months in the summer. Pinus maximartinezii is vir-
58mm long, resinous. Fascicle sheaths 78mm tually the only pine here, but a few scattered individ-
long, light brown; the outer scales deciduous, the uals of P. leiophylla var. chihuahuana have been seen.
inner scales recoiling, semi-persistent, forming a Its altitudinal range is 17502400m a.s.l. Abundant
small rosette at base of fascicle. Leaves in fascicles are various large leaved species of deciduous
of 5, very rarely 3 or 4, persisting two years, straight, Quercus, e.g. Q. macrophylla, which are bare during
lax, 711(13)cm long, 0.50.7mm wide, with entire the long dry season from September to May. Fires
margins, acute, glaucous green, in some trees green, occur regularly in the region in all vegetation types;
730 the two adaxial faces often whitish. Stomata only on it is not known whether this species is adapted well
adaxial faces. Pollen forming an elongated spike, to reseed itself after fire. Pollen dispersal is usually in
ovoid-oblong, 810mm long, yellowish. Seed cones May-June; the ovuliferous cones take 1824months
lateral, solitary on ultimate branches, on a very short to reach maturity, and perhaps longer for the seeds
peduncle or almost sessile. Mature cones broadly to ripen fully, which mostly remain in the cones.
ovoid-truncate, (15)1725(27) 1015cm when Squirrels are capable of biting off the apohyses to
the scales have parted. Seed scales parting slowly, reach the seeds and probably store them. They, and
usually insufficiently wide to release the seeds, very probably also birds, may play a crucial role in effec-
thick woody, rigid, obtrullate, up to 50mm wide tive seed dispersal, but this has not been investigated
below the apophysis, of similar shape around the to date.
cone but differentiating from base to apex, with
deep seed cavities on adaxial side. Apophyses very Conservation
prominent, 3550 2035mm, rhombic-pyramidal
at mid-cone, usually straight, transversely keeled, The species is considered Endangered due to fire and
with angular upper margin, those on the proximal grazing hazards resulting in few seedlings that suc-
and distal scales narrowly conical, often curved, dull ceed in establishing themselves. Intensive harvest-
light brown or reddish brown, often resinous. Umbo ing of cones and seeds may also diminish its chances
dorsal, obtuse-triangular or rhombic-pyramidal, of reproduction. Although there is an awareness
sometimes with a minute prickle, concolorous or amongst botanists and foresters in Mexico of its
grey-brown. Seeds oblong or ovoid-oblong, slightly importance, current in situ protection (fire pre-
flattened, 2028 (8)1012 710mm, wingless vention and fighting) seems inadequate to ensure
when the seed is free; integument ca. 2mm thick, long term preservation of this interesting narrow
very hard. Seedlings large, with 1824 cotyledons; endemic. Seed collections have been made in recent
juvenile leaves curved, flattened, ca. 8cm long, sil- years to ensure ex situ conservation programmes.
very-blue, persisting well beyond the development The land on which most of these pines grow is pri-
of adult leaves, sometimes up to 20 years. vately owned by villagers, who have an interest in the
seed harvests as well as cattle grazing.
Distribution IUCN: EN [B1ab (ii, iii, v), B2ab (ii, iii, v)]
nurseries; its horticultural merits could be greater Mature cones narrowly ovoid to ovoid-attenuate
than that since it is not too difficult to grow from when closed, more or less ovoid, slightly curved, with
seed in the nursery. Young trees retain an attractive an obliquely flattened base when opened, (4)5
blue juvenile foliage for several years. 10(12) (3)48cm when open. Seed scales thin
woody, flexible, oblong, straight, spreading wide or
often reflexed. Apophysis flattened or slightly raised
Pinus maximinoi H. E. Moore, Baileya 14: 8. 1966. and then transversely keeled, 815mm wide, irregu-
Pinus tenuifolia Benth., Pl. Hartweg.: 92. 1842, non larly rhombic to pentagonal in outline, variably light
Salisb. (1796); Pinus douglasiana Martnez var. brown. Umbo dorsal, raised, curved, transverse-
maximinoi (H. E. Moore) Silba, Phytologia 68: 50. rhombic in outline. Seeds obliquely ovoid, slightly
1990. Type: Guatemala: [In montibus para ruptis flattened, 46 34mm, ochraceous or dark brown, 731
que Canales dictis, nec non ad pagum Chinanta with or without dark spots. Seed wings oblong,
prope Guatemala, et in summo jugo Choacas prope with a straight side, widest near the middle, 1322
Salama.], C. T. Hartweg 620 (holotype K). 48mm, yellowish brown, translucent.
Etymology Distribution
This species was named after the Mexican botanist Mexico: Sinaloa, Jalisco, Michoacn, Mxico,
Maximino Martnez (18881964), who studied the Hidalgo, Tlaxcala, Puebla, Veracruz, Guerrero,
pines of Mexico. Oaxaca and Chiapas; Guatemala; Honduras; El
Salvador and NW Nicaragua.
Vernacular names TDWG codes: 79 MXC-ME MXC-PU MXC-TL MXE-
HI MXG-VC MXN-SI MXS-CL MXS-GR MXS-JA MXS-
Thin-leaf pine; ocote, pino cans (Mexico, Honduras) MI MXS-NA MXS-OA MXT-CI 80 ELS GUA HON NIC
Description Ecology
Trees to 2040(50) m tall, d.b.h. to 7090 Pinus maximinoi is a species with a wide ecologi-
(100+)cm; trunk monopodial, straight. Bark thick cal amplitude, occurring from wet subtropical for-
on lower part of trunk, with relatively smooth, est, where it is a gap pioneer, well up into the cooler
longitudinal plates and deep longitudinal fissures, cloud forests on high mountains in Mesoamerica. In
grey-brown. Branches spreading or ascending, Mexico it also occupies drier sites as a constituent of
forming a pyramidal crown in young trees and an pine or oak-pine forest or woodland. Its altitudinal
open or dense, rounded crown in mature trees. range is great: (450)6002800m, with an optimum
Shoots uni-nodal, green or light brown, rarely glau- at 9001800m a.s.l. In the NW of its range the spe-
cous. Cataphylls subulate-caudate, soon reflexed, cies occurs between 15002800m. It occurs on a
1015 mm long. Vegetative buds ovoid-conical; variety of soils under various climatic conditions; in
terminal bud 1520mm long; lateral buds smaller, Mesoamerica annual precipitation ranges from ca.
not resinous. Fascicle sheaths 1525(30)mm long, 9002500mm, with the wettest conditions on the
persistent, (lustrous) grey-brown to grey. Leaves in Atlantic and Pacific slopes of the mountains. Under
fascicles of 5, rarely 4 or 6, persisting 22.5 years, slen- these conditions it occurs frequently with Pinus
der, lax, drooping, sometimes pendulous, 2035cm tecunumanii and Liquidambar styraciflua, at lower
long, 0.61.0(1.1)mm wide, with serrulate margins, altitudes with P. oocarpa. In secondary broad-leaved
acute, (yellowish) light green to glaucous green. forest other pines may join: P. devoniana, P. pseu-
Stomata on all faces of leaves. Pollen cones densely dostrobus, and pine woodland may prevail under a
clustered, cylindrical, 34cm 58mm when full regime of grazing or burning, with the undergrowth
grown, light pinkish brown, turning darker. Seed dominated by grasses or the fern Pteridium aqui-
cones subterminal, solitary or in pairs on distinct, linum. In Central Mexico, it grows at the higher
stout, curved peduncles which fall with cones. sites with Abies religiosa, P. ayacahuite, P. patula,
Ecology
Pinus monophylla Torr. & Frm., in Frmont, Rep.
Pinus merkusii occurs in mountainous regions and Exped. Rocky Mts.: 319, t. 4. 1845, [monophyllus].
forms more or less open pine woods or pine savan- Pinus cembroides Zucc. subsp. monophylla (Torr. &
nas influenced by periodic grass fires. This ecosystem Frm.) E. Murray, Kalmia 12: 22. 1982. Type: USA: 733
is much influenced by man and may even have been California, [collected in northern California],
created by people over thousands of years of occupa- J. C. Frmont 367 (holotype NY). Fig. 231, 232
tion. Like its vicariant species on the SE Asian main-
land, P. latteri, a grass stage is reported to occur as Pinus edulis Engelm. var. fallax Little, Phytologia
an adaptation to these grass fires in the seedling to 17: 331. 1968; Pinus californiarum D. K. Bailey subsp.
sapling stages of its development. In Sumatera, this fallax (Little) D. K. Bailey, Notes Roy. Bot. Gard.
is the only pine that crosses the equator into the Edinburgh 44: 279. 1987; Pinus monophylla Torr.
southern hemisphere. & Frm. var. fallax (Little) Silba, Phytologia 68: 54.
1990; Pinus fallax (Little) Businsk, Acta Pruhon.
Conservation 88: 11. 2008.
Pinus californiarum D. K. Bailey, Notes Roy. Bot.
Pinus merkusii is threatened by over-exploitation, Gard. Edinburgh 44: 278. 1987; Pinus monophylla
habitat degradation and overgrazing. Around Lake Torr. & Frm. var. californiarum (D. K. Bailey) Silba,
Toba in northern Sumatera historical decline has Phytologia 68: 54. 1990.
fragmented the population and exploitation has pro-
gressed to the NE from there. In the Philippines the Etymology
much smaller populations are now also fragmented.
IUCN: VU [B2ab (ii, iii, v)] The species epithet monophylla refers to the single
leaf (needle) in a fascicle.
Uses
Vernacular names
Merkuss pine has been extensively planted through-
out Indonesia (where it is only indigenous in north- Singleleaf Pinyon pine
ern Sumatera) by the Dutch in colonial times.
Indonesian foresters have continued this practice as Description
it is the countrys most important producer of pine
resin. Young planted trees are better for tapping Trees or a large shrubs to 1520m tall, d.b.h. to
than old growth trees in natural stands. Indonesia 4050cm. Trunk monopodial, usually short, low
is a major producer of turpentines distilled from branched, straight or contorted. Bark thick, rough
this resin. In the Philippines, this species is tapped and scaly, with shallow, fissures, exfoliating in small
together with P. kesiya, which is indigenous on plates; outer bark reddish brown to grey. Branches
these islands, but not in Indonesia. When trees have spreading or ascending, forming a wide spreading,
grown beyond good yield of resin, their wood is har- rounded but irregular, open crown often extending
vested for the pulp industry to manufacture paper, low above the ground. Shoots short, stout, orange-
a process which allows final extraction of the resin yellow, turning grey after 12 years. Cataphylls short,
in the wood. If well managed, these plantations are triangular, rigid and spreading. Vegetative buds
a renewable resource and can assist in the preserva- ovoid-conical, acute; terminal bud 1015 57mm;
tion of the natural stands of P. merkusii. The wood lateral buds smaller, (slightly) resinous. Fascicle
sheaths of young fascicles to 11mm long, soon dis- 3000m a.s.l. The undergrowth is dominated by sage-
integrating in recoiling but almost simultaneously brush (Seriphidium spp. [Artemisia]) and numerous
deciduous scales of which only the basal parts per- other xerophytic shrubs are common. In southern
sist (not forming rosettes). Leaves in fascicles of 1, California and Baja California, P. monophylla is
rarely 2, persisting 48 years, rigid, curved at least locally common in, or just above, a high chaparral
near base, (2)2.56 cm long, 1.22.2(2.5) mm zone but does not form extensive Pinyon-Juniper
wide, in rare fascicles of 2 with entire margins, acu- woodland. Other common pines here are P. quadri-
minate-pungent, usually lustrous, green, grey-green folia and P. jeffreyi, the latter species extending to
or glaucous green. Stomata around the leaves, with higher altitude. Juniperus californica and Quercus
1225 lines in grooves. Pollen cones ovoid-globose turbinella commonly occur with P. monophylla in
734 to short cylindrical, up to 10mm long when shed- areas with granite rock outcrops and on steep slopes.
ding pollen, initially purplish red, then yellowish. Annual precipitation is low to moderate from 200
Seed cones sub-terminal, solitary or in whorls of 600mm, highly variable, in California concentrated
24 on slender, 510mm long peduncles which are in the winter and part of it as snow at higher alti-
deciduous with cones. Mature cones ovoid-globose tudes. Severe frost can occur in the Great Basin,
to globose when closed, irregular when opened, which has a continental climate.
often somewhat wider than long, 46 4.57cm
when open. Seed scales parting widely, spreading Conservation
but not reflexed, moveable, irregular, concavo-con-
vex, with 12 deep seed. Apophysis thick woody, IUCN: LC
raised, pyramidal or obtuse conical, transversely
keeled, recurved or straight, lustrous, ochraceous to Uses
yellowish brown, often resinous. Umbo dorsal, flat
or obtuse-pyramidal, rhombic in outline, centrally Due to its irregular shape and slow growth, this tree
indented. Seeds obliquely obovoid, 1318 812mm, is not used for lumber; locally it is used as firewood.
integument thin (0.30.5mm), greyish brown to The edible seeds are locally harvested and sold on
grey. Megagametophyte (endosperm) white. Seed village markets or alongside the highways. In coun-
wings absent from the free seed. tries with hot summers this would be an excellent
small pine for amenity planting, but it is little used
Distribution and only present in a few collections. In the USA,
where pines are grown for Christmas trees, it can
USA: Arizona, SW New Mexico, California, S Idaho, be put to this use, but it is slow growing and needs
Nevada, W Colorado, Utah; NW Mexico: Baja much clipping to attain the desirable shape.
California Norte.
TDWG codes: 73 COL IDA 76 ARI CAL NEV UTA 77
NWM 79 MXN-BC Pinus montezumae Lamb., Descr. Pinus, ed. 8, 1:
39, t. 22. 1832.
Ecology
Etymology
This species of pinyon pine is common on the
dry mountain slopes of the Great Basin. It is the Montezuma II was the Aztec emperor killed in a riot
major component of the extensive Pinyon-Juniper in Tenochtitln (Mexico City) in 1520 shortly before
woodland in this area; the most common juniper the Spanish conquest.
is Juniperus osteosperma, in the NW replaced by
J. occidentalis. At higher altitudes P. ponderosa can Vernacular names
be mixed in as an indicator of the transition to tall
pine forests; in the White Mountains of California Montezuma pine, Rough-branched Mexican pine;
P. monophylla has been found with P. longaeva ocote blanco, pino de Montezuma, pino real
at 3000m. Its altitudinal range is from 950m to (Mexico).
Pinus montezumae Lamb. var. montezumae. Type: TDWG codes: 79 MXC-PU MXE-QU MXG-VC MXS-
Mexico: Veracruz, [inter Cruz Blanca et Jalacingo, GR MXS-JA MXS-MI MXS-NA MXS-OA
reg. frig. Nov. 28], C. J. W. Schiede & F. Deppe s.n.
(lectotype BM). Conservation
this pathogen occurs. It is present in a few arboreta clusters near base of new shoots, spirally arranged,
and landscape parks, but uncommonly used in hor- ovoid-oblong to cylindrical, 1.52.5cm long, yellow
ticulture and only a few cultivars are known from turning light brown. Seed cones in whorls of 34 on
this species. short, stout, curved peduncles, erect at first, becom-
ing pendulous when growing, (narrowly) ovoid-
ellipsoid, 611cm long, green or glaucous, very
Pinus morrisonicola Hayata, Gard. Chron., ser. 3, resinous, with opened scales to 56cm wide, becom-
43: 194. 1908. Pinus parviflora Siebold & Zucc. var. ing more ovoid. Seed scales thin woody, somewhat
morrisonicola (Hayata) C. L. Wu, Acta Phytotax. flexible, with basal smaller scales mostly recurved
Sin. 5 (3): 141. 1956. Type: Taiwan: [Shokakulin], and larger scales nearly straight, 33.5cm long,
738 C. Owatari s.n., 21 Jan 1898 (holotype TI). 1.52cm wide, cuneate from their base; apophyses
rhombic, middle portion thickened, longitudinally
Pinus formosana Hayata, J. Linn. Soc., Bot. 38: 297. grooved, becoming lustrous brown; umbo termi-
1908. nal, obtuse, slightly upturned. Seeds ellipsoid-ovoid
Pinus uyematsui Hayata, Icon. Pl. Formos. 3: 192, t. or narrowly ovoid, 710mm long, 56mm wide;
35. 1913. wing 1520mm long, 58mm wide, light brown.
Pinus hayatana Businsk, Willdenowia 34 (1): 245.
2004. [putative hybrid between P. uyematsui and Distribution
P. morrisonicola]
Taiwan.
Etymology TDWG codes: 38 TAI
of provenance were kept with the plants in tree or excentric, armed with a minute, often breaking
nurseries. prickle. Seeds small, 34mm long; wing articulate,
1013mm long.
Pinus mugo Turra, Giorn. Italia Sci. Nat. 1: 152. Taxonomic notes
1764.
Pinus mugo is probably the species of pine with the
Etymology longest list of synonyms. Central and East European
botanists in particular have described the observed
The species epithet uses the Italian vernacular name variation in this pine as distinct species, subspecies,
for this pine. or varieties in almost every conceivable nomencla- 739
tural combination. Despite several efforts at dis-
Vernacular names tentangling the taxonomy and nomenclature, a
consensus may still elude us as some recent papers
Dwarf mountain pine, Mountain pine, Mugo pine; seem to attest. A full synonymy following what was
Bergkiefer (German); pin de montagne (French); then in my view the most reasonable taxonomy
mugo (Italian); kosodrzewina (Polish). by 2000 has been given in my World Checklist &
Bibliography of Conifers (Farjon, 1998, [2001]), sev-
Description eral new names had to be added since and it does not
look as if it will end there yet. The only subspecies
Shrubs to 5m tall, main branches ascending to erect, recognized here is subsp. rotundata, which is usu-
sometimes a small erect tree. Bark breaking into ally a small erect tree. The distinct asymmetric cones
small, rectangular plates and sometimes curling with prominent apophyses on the exposed side have
scales on larger stems, grey. Foliage branches slen- been found in some populations of shrubby forms
der, smooth with prominent ribs, glabrous, at first and are not restricted to the tree forms of the western
light greenish brown, later reddish brown to grey- Alps and Pyrenees. The seed cones of P. mugo appear
ish black. Buds cylindrical, acute, strongly resinous, to be highly variable and too much taxonomic sig-
1015mm long. Leaves in fascicles of 2, rarely 3, held nificance has been given to these variations.
in 1018mm long, later much shorter, basal fascicle
sheaths, densely set on branches, directed forward, Distribution
persisting (2)49(10) years, (2.3)37(8) cm
long, slightly curved and rigid, often twisted, 1.5 Europe, in mountains from S France to Bulgaria and
2.2mm wide, dark or light green; margins minutely Romania.
serrulate; apex more or less pungent; stomata in TDWG codes: 11 AUT-AU CZE-CZ CZE-SL GER
fine lines on all sides. Pollen cones clusterd at base HUN POL SWI 12 FRA-FR 13 ALB BUL ITA-IT
of new shoots, spirally arranged, short cylindrical, ROM YUG-BH YUG-CR YUG-KO YUG-MN YUG-SE
1012mm long, 5mm wide, yellow or red. Seed YUG-SL 14 UKR-UK
cones solitary or in whorls of 23 on short peduncles
below shoot apex, or sometimes more in a cluster Ecology
near its base and subsessile, patent or more or less
reflexed, falling at maturity or persisting up to 4 There are two principal growth forms of Pinus mugo,
years, (obliquely) ovoid, 26cm long, 24.5cm wide commonly recognised as distinct taxa: a shrub-
when opened, with an asymmetrical or symmetrical, like, sometimes nearly decumbent form (subsp.
flattened base. Seed scales thin woody, rigid, oblong, mugo) and an erect tree (subsp. rotundata), which
spreading wide, reddish or blackish brown; apoph- occupy different habitats. The shrubby form grows
yses nearly flat to prominently raised, gibbous or on mountain slopes and ridges from about 600m
transversely keeled, on the outer, sun-exposed side to 2700m a.s.l. in the mountain ranges of Europe
of cone sometimes larger and recurved or longitu- most exposed to storms associated with depres-
dinally ridged, lustrous yellowish or reddish brown; sion systems in the North Atlantic. Especially in
umbo dorsal, small, depressed, flat or conical, central the Carpathians, it forms dense mat-like thickets
above montane forests dominated by Fagus or Picea; scales usually more or less equally developed around
in the western Alps the upright form (subspecies) cone, but variable.
dominates on nutrient poor slopes. Pinus mugo in
the eastern Alps may have replaced original Larch- Distribution
Arolla pine woods which were disturbed by human
activities and grazing of their animals. The species Europe: Alps (rare in W Alps), Jura, Vosges,
often occurs on dolomite limestone, but is in fact Erzgebirge, Bhmerwald, Sudeten, Carpathians,
indifferent to soil type; this prevalence probably has Rhodope Mts., Dinaric Alps, central Apennines.
historical reasons (Ellenberg, 1988). While upright TDWG codes: 11 AUT-AU CZE-CZ CZE-SL GER POL
stands of P. mugo subsp. rotundata can be fairly SWI 12 FRA-FR 13 ALB BUL ITA-IT ROM YUG-BH
740 rich plant communities, the associated species with YUG-CR YUG-KO YUG-MA YUG-MN YUG-SE
the decumbent subsp. mugo are much fewer due to YUG-SL 14 UKR-UK
harsh environmental conditions, such as exposure
and long-lasting snow cover. Conservation
Uses IUCN: LC
2 subspecies are recognized: Small, erect trees, rarely erect shrubs. Seed cones
asymmetrical; apophyses of outer (sun-exposed)
seed scales enlarged, curved, longitudinally ridged
Pinus mugo Turra subsp. mugo. Types: Italy: Alto or keeled.
Adige, Vicenza, Monte Baldo, J. F. Sguier s.n.
(lectotype [NIMES]) & J. B. Saint-Lager s.n. Distribution
(epitype G).
Europe, Alps, Erzgebirge, Bhmerwald, Sudeten,
Description NW Carpathians.
TDWG codes: 11 AUT-AU CZE-CZ CZE-SL GER HUN
Shrubs to 5m tall, sometimes decumbent. Seed POL SWI 12 FRA-FR 13 ITA-IT
cones more or less symmetrical; apophyses of seed
not known in horticulture. It should be suitable ovoid or slightly curved ovoid, usually light brown
to Mediterranean-type climate regions and other or pale grey-brown, rarely darker. Seed scales thin
dry, warm temperate areas in Europe, the USA, woody, rigid, narrowly oblong; apophyses slightly
Australia, and South Africa. raised, transversely keeled, light yellowish brown,
more or less lustrous; umbo dorsal, small, usually
unarmed or with a minute, deciduous prickle. Seeds
Pinus nigra J. F. Arnold, Reise Mariazell: 8, t. s.n. obovoid, slightly flattened, (4)68mm long, grey
1785. or mottled darker; wing obliquely oblong, 1525mm
long, pale light brown.
Etymology
744 Taxonomic notes
The species epithet means black, this probably refers
to the bark which is much darker than that of P. syl- Few other species have a taxonomic history as con-
vestris, with which it is often found together. voluted as Pinus nigra, with numerous taxa at all
ranks from species down to forma decribed and
Vernacular names named, often in a three tiered system with varieties
within subspecies within species (for a fairly com-
Austrian pine, Black pine; pino nigro (Spanish); pin prehensive list of names and their synonyms with
noir (French); Schwarzfhre (German); Karaam references see Farjon, 1998, [2001]). It is almost
(Turkish) impossible to list all the synonyms as many were
published in regional Floras or periodicals that
Description have not even reached the major botanical librar-
ies. Especially in southern and southeastern Europe
Trees to 40m tall (50m in Corsica), max. d.b.h. a tendency to split this species on perceived differ-
1.89m (a tree in Corsica), monopodial but some- ences of all kinds, including numbers of resin ducts
times with multiple stems from a low basal trunk. in needles, is still prevalent (see e.g. Vidacovi,
Bark becoming thick and breaking into scaly ridges 1991). While it is appropriate to recognize a few
or large plates, separated by irregular, dark fissures, infraspecific taxa in a species with a variable mor-
grey with pinkish or purplish hue, dark grey-brown phology and a wide but disjunct distribution,
or nearly black. Branches spreading and ascending, extreme splitting would amount to a refutation that
sometimes heavy, forming broadly conical to domed a species after all is made up of variable individu-
crowns. Foliage branches stout, rough with pulvini als and ditto populations. A conservative treatment
from fallen leaf fascicles, glabrous, new shoots yel- is favoured here, largely based on the treatment in
lowish green, becoming light orange-brown to red- Flora Europaea 1 (1993). Even among the five sub-
brown. Buds ovoid to oblong-conical, sharply acute, species recognized here, the distinctions are often
non-resinous or resinous; cataphylls brown, thin, not as clear-cut as is claimed by those who are thor-
with papery, grey fringes. Leaves in fascicles of 2, oughly convinced of them.
held by a persistent, 1012mm long basal sheath,
remaining 23 years on branchlets, straight or more Distribution
often curved, rigid or flexible, (4)816(18)cm
long, sometimes twisted, 12mm wide, light green SW, S and SE Europe; N Algeria; N Morocco; Cyprus;
or dark green; margins minutely serrate; apex acute; Turkey; from the Krym [Crimea] in Ukraine along
stomata in fine lines on all faces. Pollen cones clus- the Black Sea coast eastwards to Krasnodar in the
tered near base of new shoots, spirally arranged, Caucasus.
ovoid-conical to short cylindrical, 1.52.5cm long, TDWG codes: 11 AUT-AU 12 COR FRA-FR SPA-AN
0.50.7mm wide, yellow. Seed cones solitary or in SPA-SP 13 ALB BUL GRC ITA-IT KRI ROM SIC-SI
whorls of 25 on short peduncles, falling shortly TUE YUG-BH YUG-CR YUG-KO YUG-MA YUG-MN
after seed dispersal, ovoid-conical when closed, YUG-SE YUG-SL 14 KRY 20 ALG MOR-MO 33 NCS-KR
(3.5)510(12)cm long, 24cm wide, opening to 34 CYP EAI TUR
layers and 1011 resin ducts; seed cones small, (3)4 Pinus nigra J. F. Arnold subsp. pallasiana (Lamb.)
6(7)cm long. Holmboe var. pallasiana, [Stud. Veg. Cyprus]
Bergens Mus. Skr., ser. 2, 1 (2): 29. 1914. Pinus pal-
Distribution lasiana Lamb., Descr. Pinus 2: 1, t. 1. 1824. Type:
Illustration in Lambert, Descr. Pinus 2: t. 1. 1824
SE Europe: Croatia. (lectotype, designated here).
TDWG codes: 13 YUG-CR
Pinus laricio Poir. var. caramanica Loudon, Arbor.
Conservation Frut. Brit. 4: 2201. 1838; Pinus nigra J. F. Arnold
var. caramanica (Loudon) Rehd., Man. Cult. Trees:
746 This subspecies of Pinus nigra has a limited distri- 61. 1927; Pinus nigra J. F. Arnold subsp. caramanica
bution along the coast and on some islands in the (Loudon) Businsk, Acta Pruhoniciana 68: 22. 1999;
Adriatic Sea and is mainly threatened by habitat Pinus pallasiana Lamb. subsp. caramanica (Loudon)
degradation. Many areas are seriously overgrazed Chrtek & B. Slavik, Flora Mediterranea 10: 236. 2000.
especially by goats, preventing regeneration. Pinus nigra J. F. Arnold var. yaltirikiana C. U.
IUCN: EN [B1ab (iii, v), B2ab (ii, v)] Alptekin, Istanbul Univ. Orman Fak. Dergisi, ser. A,
36 (2): 147. 1987.
Pinus nigra J. F. Arnold var. columnaris-pendula
Pinus nigra J. F. Arnold subsp. laricio (Poir.) Maire, Boydak, Karaca Arbor. Mag. 6 (1): 17. 2001.
Bull. Soc. Hist. Nat. Afrique N. 19: 66. 1928. Pinus
laricio Poir., in Lamarck, Encycl. 5: 839. 1804, non Description
Savi (1798. Type not designated. Fig. 236
Trees to 40m tall, commonly (in cultivation?)
Pinus laricio Poir. var. calabrica Loudon, Arbor. Frut. forked into 2 or more ascending trunks; bark light
Brit. 4: 2201. 1838; Pinus nigra J. F. Arnold var. cal- grey, with deep, longitudinal, dark fissures and often
abrica (Loudon) C. K. Schneid., in Silva-Tarouca, very large, irregular, scaly plates. Foliage branches
Uns. Freil.-Nadelhlzer: 261. 1913; Pinus nigra orange-brown; leaves rigid, straight or curved,
J. F. Arnold subsp. calabrica (Loudon) E. Murray, (8)1217cm long, light green, with 25 hypodermal
Kalmia 13: 23. 1983; Pinus laricio Poir. subsp. cal- cell layers and 69 resin ducts.
abrica (Loudon) Cesca & Peruzzi, Caryologia 55 (1):
24. 2002. Distribution
Distribution
Pinus nigra J. F. Arnold subsp. pallasiana (Lamb.)
Mediterranean: France (Corsica); Italy (S Apennines, Holmboe var. fastigiata Businsk, Acta Pruhon. 88:
Sicily). 8. 2008.
TDWG codes: 12 COR 13 ITA-IT
Description
Conservation
A fastigiate branching growth form of Pinus nigra
IUCN: LC subsp. pallasiana.
Turkey: Ktahya, near Dulkadir village. Pinus occidentalis Sw. var. baorucoensis Silba, 747
Phytologia 58: 368.
3.56.5cm when open. Seed scales oblong, straight Darrow & Zanoni (1991) it has been depleted from
or recurved, thin woody. Apophysis slightly raised, an estimated 3million ha of primeval more or less
transversely keeled, rhombic to pentagonal in out- pure pine forests to perhaps less than 5% of that
line, on the proximal scales more or less gibbous, area, but accurate estimates of even the present
(lustrous) dark brown, radially striate, weathering forest extent are lacking. Protection in the Dominican
dull grey. Umbo dorsal, raised and often curved, Republic is inadequate, but existent, contrary to the
usually armed with an inflexed, 23mm long spine. virtually uncontrolled situation in Haiti.
Seeds obliquely obovoid, flattened, 56 34mm, IUCN: EN [A2a, c, d; B2ab (ii, iii, v)]
light, mottled grey-brown. Seed wings obliquely
ovate or oblong, 1218 46mm, ochraceous with Uses
748 black or grey tinge or stripes.
Cuban pine (Hispaniolan pine) is an important
Distribution timber tree on its native island Hispaniola, where
despite intensive exploitation it is still common. Its
Hispaniola: Dominican Republic; Haiti. wood has good qualities comparable to those found
TDWG codes: 81 DOM HAI-HA in the more widespread species P. caribaea and is
used as round wood for transmission poles, fence
Ecology posts, construction timber, crates, boxes, and made
into wood pulp for particleboard as well as paper.
This species occurs in diverse habitats from the There is limited resin tapping for local use only. It
lowlands at about 200m a.s.l. to the highest moun- is a tropical pine and even those trees at the highest
tain ridges (Pico Duarte and Pico La Pelona) on the altitudes in the Dominican Republic are not likely
island at almost 3200m. The more extensive and to yield progeny that can be grown successfully in
pure stands occur from 9002700m, but in more cool temperate climates. Growing pines in green-
accessible areas these are much depleted. Soils are houses is problematic: unlike many other subtropi-
either derived from limestone at lower altitudes, or cal or tropical conifers they do not produce strong
more acid, clay-like and shallow in the Cordillera wood to hold the tree upright without the influence
Central. Pinus occidentalis consequently is found of wind that moves the stems during secondary
in a variety of vegetation types, mostly occupying growth.
the shallow, nutrient-poor soils and rock outcrops,
where it may occur in open or dense, pure stands or
mixed with various broad-leaved trees and shrubs. Pinus oocarpa Schiede ex Schltdl., Linnaea 12:
In disturbed (grazed) areas Pteridium aquilinum can 491. 1838. Type: Mexico: Michoacan, Aguililla, Las
dominate the ground cover; in frequently burnt areas Playitas Forest Station, along the track to Caas,
grasses (e.g. Danthonia domingensis, Andropogon B. T. Styles 36 (neotype FHO).
spp.) and again Pteridium replace shrubs and small
trees. Annual precipitation varies greatly with expo- Pinus oocarpa Schiede ex Schltdl. var. manzanoi
sition, but ranges between 12001600mm where Martnez, Anales Inst. Biol. Univ. Nac. Mxico 11:
most pine forests occur, it exceeds 2300mm in the N 70. 1940.
and E of the Cordillera Central. There is a 35month
dry season during winter, which may bring frost, Etymology
but rarely snow, at the higher altitudes above
16001800m. The species epithet (Greek: oos = egg, carpos = fruit)
refers to the shape of closed seed cones.
Conservation
Vernacular names
Being the only species of Pinus on Hispaniola and
formerly abundant over much of the island, it has Egg-cone pine; ocote chino, pino chino, pino colo-
been heavily exploited for timber. According to rado (Spanish)
markets. The wood is of better quality than that lines on all surfaces. Pollen cones clustered at base of
of P. caribaea, it is stronger and less resinous. It is new shoots, spirally arranged, spreading out radially,
much used for sawn timber, especially applied to cylindrical, 47(8)cm long, 0.81.5cm wide, pur-
light construction like carpentry and joinery and plish red, turning red-brown. Seed cones solitary or
floors in houses. Courser uses are railway sleepers in pairs near ends of foliage branches, sessile or very
and beams or transmission poles. The species is short pedunculate, soon falling after seed dispersal,
also widely tapped for resin, but only for a few years often leaving a rosette of basal scales on branch, (15
before felling. This species has been introduced to )2025cm long, symmetric, narrowly ovoid when
many subtropical and tropical countries as a plan- closed, broadly ovoid-cylindric when open, with
tation forestry tree, it is perhaps most successful in a flat base. Seed scales oblong, thin woody, rigid,
750 W Africa, South Africa, and in Brazil, Colombia and recurved near base of cone, elsewhere spreading
Argentina. In horticulture it is very rare and limited wide, dull brown; apophyses slightly raised, trans-
to a few botanic gardens and other tree collections in versely keeled, rhombic in outline, dull tan or brown
countries with warm climates. in colour; umbo dorsal and central, broadly triangu-
lar, obtuse, armed with a small, persistent, recurved
prickle. Seeds obovoid, slightly flattened, 912mm
Pinus palustris Mill., Gard. Dict., ed. 8: Pinus No. long, pale brown mottled with darker specks; wing
14. 1768. Type not designated. 3040mm long, 1012mm wide. Seedlings with a
definite and prolonged grass stage.
Etymology
Distribution
The species epithet means of the marsh, referring to
the habitat of the species. SE USA, from Virginia to E Texas in the Atlantic and
Gulf Coastal Plains.
Vernacular names TDWG codes: 77 TEX 78 ALA FLA GEO LOU MSI
NCA SCA VRG
Longleaf pine, Florida pine, Georgia pine
Ecology
Description
Pinus palustris grows mostly in the warm temperate
Trees to 45(47)m tall; trunk to 1.2m d.b.h., bole to subtropical coastal plain of the SE United States,
straight and columnar. Bark breaking into large, but it extends into the uplands and foothills of the
irregularly rectangular, scaly plates separated by deep southern Appalachian Mountains to about 700m
fissures, orange-brown or reddish brown. Branches a.s.l. It requires a warm, humid climate and a moist
relatively sparse, spreading and descending, forming soil, although the latter can vary from wet, poorly
an open, rounded crown. Foliage branches stout, up drained clay in swampland to thin stony soils on
to 2cm thick in leading shoots, upturned at apex, rocky mountain slopes and ridges. For the most part
prominently ridged and grooved between decurrent soils are sandy, acidic and poor in nutrients. This
pulvini, glabrous, orange-brown turning darker red- species can grow in pure stands but is often found
dish brown. Buds large; terminal bud 34.5cm long, together with other pines, like P. elliottii, P. echi-
narrowly ovoid, not resinous; cataphylls narrowly nata and P. taeda. Many broad-leaved trees (angio-
oblong, with fringed margins and recurved at apex, sperms) grow with it, too, forming mixed forests on
silvery white. Leaves in fascicles of 3, rarely 2 or 5, mesic sites with e.g. Quercus spp., Nyssa sylvatica,
held together in 2025(30)mm long sheaths, per- Liquidambar styraciflua, Cornus florida, Sassafras
sisting 2 years, 2045cm long (longest in young, vig- albidum, and Diospyros virginiana and many shrubs.
orous trees), spreading and drooping, slender and In the foothills in Alabama many oaks (Quercus
flexible, slightly twisted, ca. 1.5mm wide; margins spp.) accompany Pinus palustris, forming mixed
minutely serrulate; apex shortly acute to acuminate; pine-oak woods. This species of pine is well adapted
leaf colour bright lustrous green; stomata in fine to fires through its grass stage and succeeds quickly
from seed to take advantage of freed space and Pinus parviflora Siebold & Zucc., Fl. Japon. 2 (3):
nutrients, while it can survive the next ground fire 27, t. 115. 1842.
by resprouting from the very short stem until it has
built enough of a root system to accelerate its stem Etymology
growth dramatically and rise above the damaging
flames. The species epithet means with little flowers which
perhaps refers to an early stage of the seed cones.
Conservation
Vernacular names
Pinus palustris was once the most common pine in
the Coastal Plains, perhaps covering 25million ha. Japanese white pine; goy-matsu, himeko-matsu 751
Exploitation for timber and naval stores and conver- (Japanese)
sion to farmland and pasture reduced this by 1985
to less than 1.6million ha. This decline, although Description
its rate has slowed, is ongoing. Foresters and land-
owners prefer other species that do not delay initial Trees to 1520m tall; trunk to 1m d.b.h., usually a
growth with a grass stage and many sites originally straight, columnar bole. Bark smooth in young trees
occupied by P. palustris are now dominated by other and in upper part of the crown, becoming rough and
species. Prevention of fires increases competition scaly on trunk, shallowly grooved, grey to dark grey.
from herbs and shrubs as well as other pines, as Branches whorled in young trees, later spreading
the seedlings of P. palustris do not initially grow in and ascending; stem sometimes forked near the top,
height, unlike other pines common on the Coastal forming a conical to rounded crown, on exposed sites
Plains. Controlled burning is therefore necessary often flattened and open. Foliage branches slender,
for its successful establishment, but only P. palustris new shoots initially pubescent, soon glabrous or with
will survive this at a juvenile stage and in managed remnant hairs in grooves, grey-green turning light
pine forest this practice would exclude other pine brown. Buds small, ovoid, non-resinous or slightly
species. resinous; cataphylls free at apex, light orange-brown.
IUCN: EN (A2c, d, e) Leaves crowded in dense tufts towards end of shoots,
persisting 35 years, in fascicles of 5, held by decidu-
Uses ous sheaths with thin scales falling in the second
year, curved and slightly twisted, (3)47(8)cm
Longleaf pine is an important species provid- long, slender and flexible, 0.71mm wide, trian-
ing good quality timber as well as naval stores gular in cross-section, often twisted, green to glau-
derived from its resin. It was heavily exploited since cous green; stomata in 34 prominent white lines
Europeans settled in the Coastal Plains and served a on the two adaxial surfaces; margins minutely ser-
major forest industry in the region. Its wood is used rulate; apex acute. Pollen cones in small clusters
for sawlogs, stage flooring, plywood, pulpwood and near base of new shoots, spirally arranged, ovoid-
produces poles, fence posts, and piling as it makes oblong to cylindrical, 1.52.5cm long, yellow turn-
straight stems largely free of branches when grown ing light brown. Seed cones in whorls of 48(10) on
in closed stands. Turpentine and other chemicals branches, persistent, sessile or very short peduncu-
can be distilled from the chipped wood and even late, spreading, when still growing ovoid or subglo-
stumps of trees are pulled from the soil for this pur- bose, green or glaucous, very resinous, with opened
pose. Longleaf pine is not planted for forestry out- scales 47cm long and to 4cm wide, becoming more
side its natural range and it is relatively rare as an ovoid or irregular. Seed scales few, thin woody, flex-
amenity tree, mainly confined to coastal areas in ible, with larger scales straight or curved, but basal
the SE of the USA. The species is somewhat difficult smaller scales not recurved; apophyses variously
to grow through its grass stage to the sapling stage shaped, strongly concavo-convex (cupped) or more
and beyond and needs a protected location in a mild or less flat, incurved, becoming dull brown; umbo
climate. terminal, obtuse. Seeds ellipsoid-ovoid or obovoid,
710mm long, 56mm wide; wing to 12mm long, is inhibited and plants are slowly forced by various
or rudimentary. techniques into shapes that appear to imitate ancient
trees growing on exposed mountains in a miniature
Taxonomic notes fashion. In Western horticulture, the species is also
popular and a substantial number of cultivars has
Pinus parviflora is closely related to a group of pines been produced, predominantly dwarf forms with
that includes P. fenzeliana (syn. P. kwangtungen- strongly glaucous foliage and some with variegated
sis), P. dalatensis, P. wangii, and P. morrisonicola, needles. Naturally growing trees of the species are
but occurs at a considerable distance from these in relatively rare and mostly confined to collections in
Japan. One variety is commonly recognized: P. par- arboreta. Most plants in the trade under the species
752 viflora var. pentaphylla (Mayr) A. Henry (syn. P. pen- name are derived from Japanese garden selections
taphylla Mayr), with a more or less well developed (cultivars without a name).
seed wing and flatter, less cup-like seed cone scales.
Wingless seeds and hollowed, cup-like scales that 2 varieties are recognized:
accommodate them are not independent characters
but form an adaptive complex to prevent the seeds
from falling where the seedlings have little chance Pinus parviflora Siebold & Zucc. var. parviflora.
of growing up under the parent trees. Persistence of Type: Japan: [locality unknown], Pinus cembra,
cones may also be part of this syndrome, depending P. F. von Siebold s.n. (lectotype L, acc. no.
on the feeding habits of the bird species that feed on 901.138394).
them and (presumably) play a role in their dispersal.
Description
Distribution
Seed scales cup-like in shape, with deep seed cavi-
Japan: Hokkaido, Honshu, Kyushu, Shikoku; South ties. Wings of seeds 37mm long, 68mm wide,
Korea, Ullung-do (island). rudimentary (shorter than the seed body).
TDWG codes: 38 JAP-HK JAP-HN JAP-KY JAP-SH
KOR-SK Distribution
other countries where it has been introduced for Pinus patula Schiede ex Schltdl. & Cham. var. lon-
plantation forestry on a large scale are Colombia, gipedunculata Loock ex Martnez, Pinos Mexic., ed.
Brazil and Argentina and to a more limited extent 2: 333, f. 276280. 1948 [longepedunculata]. Pinus
India, Nepal and New Zealand. Much work has been longipedunculata (Loock ex Martnez) Businsk,
done by forest geneticists and tree growers in breed- Acta Pruhon. 88: 11. 2008. Type: Mexico: Oaxaca,
ing programmes to improve seed provenances for Sierra Benito Juarez, Rancho Benito Juarez, (Rancho
timber growing in tropical countries. The wood is Tablas), E. E. M. Loock 113a (holotype PRF).
soft and light coloured and easily worked, but can be
susceptible to blue stain without treatment. It finds Description
applications in flooring and panelling, plywood
754 and particleboard manufacture, veneers, crates and Seed cones distinctly pedunculate, peduncles up
boxes, and of course for its softness, pulp for paper; to 20mm long, curved, falling with cones after a
it is therefore less suitable for high quality furniture few years. Cones opening readily, narrowly ovoid
and tools. In horticulture it has made some progress to ovoid-attenuate when closed, (narrowly) ovoid
in recent years as it forms an attractive tree with its when open, then 58 3.54.5(5)cm. Seed scales
fine foliage and several provenances have proved to ca. 100120, (thin) woody, somewhat flexible, part-
be quite hardy. ing readily to release the seeds.
and branches, becoming thicker and breaking into similarities. Instead, P. cembra is probably more
small plates, at lower trunk longitudinally fissured, closely related to P. peuce than to P. albicaulis. A
grey-brown; inner bark purplish brown. Branches more careful second look at anatomy and morphol-
spreading more or less horizontally, relatively short ogy, avoiding strongly adaptive traits that have led
or longer and assurgent in solitary trees, poorly to convergence, may yet detect characters that sup-
self-pruning, forming a dense, conical or narrowly port the results from DNA studies, or refute them, as
conical, sometimes broadly conical crown. Foliage the case may be. It is important to include all species
branches stout; new shoots glabrous, smooth, often of section Quinquefolius and appropriate outgroup
glaucous, turning grey-green and finally grey-brown. taxa in such studies.
Buds ovoid-conical, ca. 10mm long, acute, resinous
or without resin; cataphylls brown with a narrow, Distribution 755
whitish margin. Leaves in fascicles of 5, initially held
in a deciduous, 1518mm long basal sheath, persist- SE Europe: Albania; W Bulgaria; Kosovo; Macedonia;
ing 34 years in tufts on branchlets, straight, more Montenegro; N Greece; Serbia.
or less rigid, 710(12)cm long, slender, 0.50.7mm TDWG codes: 13 ALB BUL GRC YUG-KO YUG-MA
wide; margins minutely serrulate; leaf colour grey- YUG-MN YUG-SE
green or slightly glaucous; apex acute; stomata in fine
whitish lines on all surfaces, but only a few on the Ecology
abaxial side. Pollen cones few and small, 1015mm
long, clustered in spiral arrangement at base of new Pinus peuce is a montane pine, growing between
shoots, yellow with a purple hue. Seed cones solitary 600m and 2200m above sea level and usually on
or in whorls of 24 on short, curved peduncles, ini- silicate rock types. In Albania and Serbia it is also
tially erect, green becoming red or purple, when fully found on serpentine. It can grow on a variety of
grown becoming pendulous, cylindrical, (7)815( soil types, usually poor in nutrients and acidic
20)cm long, obliquely curved at base, 23(4)cm to basic. In Grecian Macedonia it forms pure
wide when closed, opening to 45(6)cm wide, stands on gentle mountain slopes, interspersed
resinous. Seed scales thin woody, flexible, cune- with grassy glades and meadows. In most areas
ate to obovate, more or less flat; basal scales not where it occurs, it is mixed with Picea abies and/
recurved but imbricate; apophyses broadly rhom- or Abies alba / A. borisii-regis, with which it can
bic to rounded distally, slightly thickened, mostly compete due to relatively high shade tolerance.
incurved at apex, longitudinally striate or grooved,
light brown when dried; umbo terminal, obtuse, Conservation
dark grey-brown. Seeds obovoid, slightly flattened,
(5)68mm long, grey; wing adnate, 1520mm long. This species is probably a Tertiary relict that has sur-
vived severe contractions of its range due to Alpine
Taxonomic notes glaciations during the Pleistocene. Its current range
consists of two disjunct populations, one in the
The relationships of this species, the only European west centered in Albania and one in the east in W
member of Subsection Strobi in Europe according to Bulgaria. The two populations have sometimes been
the classification adopted in the first edition of the viewed as two distinct varieties, with Pinus peuce
book Pines (Farjon, 1984: 194), have recently been var. vermiculata Christ (cited in Vidakovi, 1991:
revised on the basis of molecular evidence (Liston 532 but orig. publ. not traced) in Bulgaria, but this
et al. in Mill, 2003). The division into subsections is not generally accepted as the stated differences:
Cembrae and Strobi based on cone and seed mor- length of leaves and cones, are continuous and partly
phology appear to break down, with similar results overlapping. Extensive exploitation in the past has
from both chloroplast and nuclear DNA. The wing- undoubtedly had an impact on its abundance but no
less seeds, dispersed by birds, have resulted in a spe- details of a possible reduction are known. At present,
cific type of cone (see e.g. P. albicaulis and P. cembra) this species occurs in several protected areas in the
amounting to convergent evolution. These pines Balkan mountains.
are not necessarily closely related because of these IUCN: NT
759
Plate 32. Pinus pinceana. 1. Habit of tree. 2. Branch with foliage and seed cone. 3. Leaves. 4. Seed cone.
5. Seed.
cylindrical, often irregular, 510 3.56(7)cm gardens with a Mediterranean climate or nearly
when open. Seed scales parting only slightly or equivalent.
sometimes more widely, easily moveable, thick
woody, concavo-convex, with (1)2 deep, cup-like
depressions holding the seeds. Apophysis irregular, Pinus pinea L., Sp. Pl. 2: 1000. 1753. Type:
prominently raised, transversely keeled, rhombic to Illustration: Pinus ossiculis duris, foliis longis in
pentagonal in outline, with angular or undulating Bauhin & Cherler, Hist. Pl. Univ. 1 (2): 248. 1650
upper margin, lustrous red-brown, sometimes with (lectotype). Fig. 242, 243
concentric colour rings or radially striated. Umbo
dorsal, flattened at apex, up to 5mm high, with a Etymology
760 minute prickle. Seeds obovoid, 1114 78mm,
wingless when detached from the scale, brown. The species epithet is a Latin adjective (with femi-
nine gender)meaning of pines (masc. = pineus).
Distribution
Vernacular names
Mexico: Coahuila, Hidalgo, Quertaro, San Luis
Potos, Zacatecas. Stone pine, Umbrella pine; pino domestico, pino
TDWG codes: 79 MXE-CO MXE-HI MXE-QU MXE- da pinoli (Italian); pino pionero, pino manso
SL MXE-ZA (Spanish); pinheiro manso (Portuguese); pin pignon,
pin parasol (French)
Ecology
Description
This species occurs often on calcareous slopes and
in ravines (barrancas) at altitudes between 1400 Trees to 2025m tall; trunk to 1m d.b.h., mono-
2300m a.s.l. in arid and semi-arid mountains of NE podial or soon forked into more stems. Bark scaly,
Mexico. Annual precipitation is lower than in the becoming thick on trunk, breaking into large,
Pinyon-Juniper belt (ca. 300400mm according to hard plates divided by deep, irregular, dark brown
Perry, 1991) and is concentrated in the summer. This to black fissures, orange-brown to reddish brown
species sometimes occurs with Pinus cembroides, with grey patches of flakes that remained longer.
more often with Juniperus flaccida, and usually with Branches numerous, spreading and ascending, the
numerous sclerophyllous or deciduous shrubs, e.g. higher order branches assurgent to erect, forming an
Prosopis (in arroyos), Mimosa, Karwinskia, Leucaena, umbrella-shaped, dense crown; young trees are more
Sophora, Quercus, Cercocarpus, Gochnatia, and or less conical with an erect stem. Foliage branches
Fouquieria, as well as the succulent genera Yucca, slender or stout, new shoots glabrous, rough with
Agave and Opuntia. The trees are often widely scat- pulvini of fallen leaf fascicles, pale green to yellowish
tered in this vegetation. brown, turning orange-brown. Buds ovoid-conical,
612mm long, acute, not resinous; cataphylls light
Conservation brown or red-brown, with fringed margins and
reflexed apex. Leaves in fascicles of 2 held by a short,
IUCN: LC persistent basal sheath, remaining 34 years, rigid
and spreading, 812(15)cm long, usually twisted,
Uses 1.21.8mm wide; margins minutely serrate; leaf
colour on young plants glaucous, becoming lustrous
This species is not commercially used; the seeds dark green; stomata in fine lines on all surfaces.
are edible (piones), but trees are scarce, inacces- Pollen cones short cylindrical, yellow. Seed cones
sible and far apart and usually with a low crop of solitary or sometimes in whorls of 23 on short
cones compared to P. cembroides and its close rela- peduncles, persisting several years, serotinous or
tives. It is not known in cultivation outside a few semi-serotinous, ripening in the third year, symmet-
botanic gardens and arboreta; it will be suitable in rical, broadly ovoid to subglobose, (8)913(15)cm
long, 810cm wide when closed, when finally open- The quality of the wood for sawn timber is poor,
ing up to 13cm wide near the depressed base, leav- being coarse and resinous and seldom straight for
ing some basal scales on branch when falling. Seed any substantial length. It is locally used to make
scales thick woody, rigid, with 2 deep depressions on furniture. The true economic value of this pine
adaxial side holding the seeds. Apophysis (slightly) has since ancient times mostly resided in the edi-
raised, angular in outline, convex with 46 ridges ble seeds. These are harvested while still inside the
converging on the dorsal, central, obtuse umbo, closed cones, which are pulled down with hooked
maturing from olive green to lustrous orange-brown poles; the cones open when heated. For this purpose
or red-brown, weathering grey. Seeds obovoid, lower tree crowns are favoured, which is accom-
1520(22)mm long, 810mm wide, brown; wing plished by keeping the canopy open. Seeds are a
mostly rudimentary, oblique, 310(15)mm long. delicacy in themselves, but are also used in recipies 761
ranging from pasta dishes to pastries. Millions of
Distribution kilogrammes of seeds or pine kernels are harvested
in the Mediterranean countries each year. Empty
Mediterranean Europe and Near East (doubtfully pine cones are good, hot burning fuel (bakeries), but
native in many areas of E Mediterranean). can also be sold as ornamental objects to florists. The
TDWG codes: 12 BAL COR FRA-FR FRA-MO POR resin is tapped and used for medicinal treatments,
SAR SPA-SP 13 ALB GRC ITA-IT KRI SIC-SI YUG-CR wax for violin bows, varnishes, and waterproofing.
34 CYP EAI LBS-LB TUR A green dye is made from the needles. The species is
also much valued as an ornamental tree in gardens
Ecology and parks around the Mediterranean and in other
parts of Europe with mild winters. In South Africa it
This is primarily a pine of coastal areas in the has become naturalized and is an invasive and nox-
Mediterranean, at elevations from sea level to 600m, ious species in the fynbos biome of the Cape.
on coastal dunes and flats as well as on lower slopes of
mountains and in the hills. Many present-day stands
are the result of historic plantings, some going back Pinus ponderosa Douglas ex C. Lawson, Agric.
to Roman times, and if managed well, these can have Man.: 354. 1836.
a natural understorey of maquis scrub or mixture
with smaller broad-leaved trees. Mature trees have a Etymology
thick, fire resistant bark and the massive cones take
three years to mature and are serotinous or semi- The species epithet (Latin, fem. of ponderosus)means
serotinous. Seeds are nearly wingless and dispersed ponderous or massive.
by birds (also eaten by rodents) or may scatter after
fire burned off the undergrowth and its heat assisted Vernacular names
in opening the cones. Pinus pinea is usually an emer-
gent tree above shrubs (maquis) or in low, open for- Ponderosa pine, Western Yellow pine
ests; it can also occur with Pinus halepensis and in
Quercus ilex maquis-woodland. Description
often long, with lower branches down-curved, con- P. benthamiana. American foresters treated this pine
torted or more or less pendulous, forming broad, as if it included the type of P. ponderosa. Nobody
columnar or domed, open crowns. Foliage branches seems to have bothered to ask what the type speci-
stout; new shoots 0.72cm thick, rough with pulvini men of P. ponderosa might be and where it had been
from fallen leaf fascicles, orange-brown, becoming collected; these questions have now been resolved
brown. Buds ovoid, to 2cm long; terminal bud to (Lauria, 1996). Only P. ponderosa var. scopulorum
1cm wide, all resinous; cataphylls red-brown with is here recognized as a botanical variety within the
white fringes. Leaves in fascicles of 23, persisting species; P. arizonica is treated as a distinct species
23 years, held by a ca. 1520(25)mm long, per- largely, but not exclusively, occuring in Mexico. If we
sistent basal sheath, spreading, straight or curved, include the smaller trees of the North Plateau Race
762 1525cm long, rigid or pliant, slightly twisted, in P. ponderosa var. ponderosa (as we must), as well
(1)1.21.8mm wide; margins minutely serrulate; as the spurious taxon P. washoensis and the tall trees
apex acute or acuminate; leaf colour green; fine of the Cascades Sierra Nevada, then the only mor-
lines of stomata on all surfaces. Pollen cones spirally phological difference that appears to hold up more
arranged, erect and standing out radially, 34.5cm or less consistently is the number of leaves per fas-
long when full-grown, ellipsoid-cylindric, initially cicle: predominantly 2, rarely 3 in var. ponderosa, 23
bright red or magenta, turning yellow at anthe- (with a N-S cline towards increasing frequency of
sis from yellow pollen sacs. Seed cones solitary or 3) in var. scopulorum. This is a difference that barely
rarely in pairs on short peduncles to nearly sessile, merits taxonomic recognition in conifers.
leaving some scales on branch when falling, nearly
symmetrical, ovoid-conical when closed, ovoid to Distribution
subglobose when opened and 512cm long. Seed
scales oblong, spreading wide or reflexed, thin W North America, from S British Columbia to just
woody, rigid. Apophyses slightly to prominently south of the Mexico-USA border.
raised, transversely keeled, more or less rhombic in TDWG codes: 71 BRC 73 COL IDA MNT ORE WAS
outline, dull tan or lustrous reddish brown; umbo WYO 74 NDA NEB OKL SDA 76 ARI CAL NEV UTA 77
raised (rarely flat), pyramidal, keeled, recurved and NWM TEX 79 MXE-CO MXN-SO
usually armed with a hooked prickle. Seeds ellipsoid
or obovoid, (4)57(8)mm long, brown or light Ecology
brown, often mottled darker; wing 1525mm long.
Pinus ponderosa is one of the most widely distributed
Taxonomic notes pines, with a range spanning 20 degrees of latitude
and from the lowlands to 3300m a.s.l. While in the
Pinus ponderosa is morphologically variable north and east its limits are defined by temperature
mainly in a north-south gradient from Canada to and rainfall respectively, in the south it is replaced
Mexico, but also from the Pacific coast to the Rocky by related species. The lower limit of average annual
Mountains. Its economic importance has stimulated precipitation is around 300mm, the upper limit
numerous studies into this variation, but much of 1750mm. It grows on soils derived from many rock
it has not been built on a sound taxonomic basis. types, both acidic and basic varying from around pH
It has appeared from taxonomic studies, that what 5 to pH 9, and usually with a capacity to retain mois-
David Douglas collected as P. ponderosa is called ture but well drained. Its great altitudinal range as
by foresters the North Plateau Race of that spe- well as geographic spread causes huge variations in
cies and is conspecific with P. washoensis, described temperature, both summer and winter, but to thrive
from Lake Tahoe in Nevada. The Ponderosa pine and compete well it requires ample sunshine in the
of the Cascade Range in Oregon and the western growing season. Pinus ponderosa is the first tall pine
slopes of the Sierra Nevada in California, while to appear above Pinyon-Juniper woodland in the
here included in P. ponderosa (Pacific Race of for- interior of the western USA, but it is a seral species in
esters), would, if sufficiently different and distinct the mesic mixed conifer forests of the Cascades and
as a species from P. washoensis, have to be called Sierra Nevada, where other conifers will eventually
dominate. It grows with numerous other conifers 1525mm long (seed character states are correlated
(for listings see under P. monticola and P. jeffreyi) in with cone size, not independent character states,
these mountain ranges. Its relatively high tolerance and larger in this variety because the cones are
of fire assures it a place in the succession in natural larger).
forests, but selective logging as well as fire prevention
have altered the forest composition to the disadvan- Distribution
tage of P. ponderosa in many managed forests.
Pacific W North America, from S British Columbia
Uses to S California.
TDWG codes: 71 BRC 73 IDA MNT ORE WAS 76
Ponderosa pine is one of the most important timber CAL NEV 763
trees in the USA. Its wood properties and potential
size make it ideally suitable for sawn timber used in Conservation
construction. It is used as roundwood for posts and
sawn for crates as well as made into particle board IUCN: LC
and pulp. Timber of large sizes and high grade is
sawn for light construction like window frames,
doors, stairs, flooring, sidings, panelling, veneers Pinus ponderosa Douglas ex C. Lawson var. scopu-
and for furniture, cabinetwork, boxes, and wood- lorum Engelm., in Watson, Bot. California 2: 126.
ware. In the National Forests of the USA the recre- 1880. Pinus scopulorum (Engelm.) Lemmon, Gard.
ational and ecological functions of the open, sunny & Forest 10: 183. 1897; Pinus ponderosa Douglas ex
forests of Ponderosa pine at relatively low altitudes C. Lawson subsp. scopulorum (Engelm.) E. Murray,
are increasingly recognized and on public lands Kalmia 12: 23. 1982. Type: USA: Colorado
these trees are no longer exclusively regarded as a [Colorado Territory, lat. 3941, Alpine and
source of timber. The species and its variety scopulo- Subalpine], C. C. Parry s.n. (lectotype MO).
rum are common in horticulture in parts of Europe,
but usually limited to large parks and arboreta. A Description
few cultivars have been named, but they are uncom-
mon. Introductions for forestry plantations have Leaves in fascicles of 23. Seed cones 510cm long;
been less successful due to fungal diseases, e.g. in seeds 35mm long; seed wings 1215mm long
New Zealand. (seed character states are correlated with cone size
and smaller in this variety because the cones are
2 varieties are recognized: smaller).
Distribution
Pinus ponderosa Douglas ex C. Lawson var.
ponderosa. Type: ovuliferous cone with appended USA: Rocky Mts., Black Hills; Mexico: near the bor-
label stating: Bayford Bury January 2 1849 der in Coahuila and Sonora.
(neotype W). Fig. 244 TDWG codes: 73 COL MNT WYO 74 NDA NEB OKL
SDA 76 ARI NEV UTA 77 NWM TEX 79 MXE-CO
Pinus washoensis Mason & Stockw., Madroo 8: 62. MXN-SO
1945; Pinus ponderosa Douglas ex C. Lawson subsp.
washoensis (Mason & Stockw.) E. Murray, Kalmia 12: Conservation
23. 1982.
IUCN: LC
Description
Pinus praetermissa Styles & McVaugh, Contr. Univ. Seeds obliquely ovoid, 58 34mm, blackish grey
Michigan Herb. 17: 310, f. 2. 1990. Type: Mexico: or with black dots; wings 1218 58mm, greyish
Nayarit, Jal, Cerro Juanacata, J. W. Stead & brown.
B. T. Styles 475 (holotype FHO).
Distribution
Etymology
Mexico: Durango, Jalisco, Nayarit, Sinaloa.
The species epithet means overlooked or neglected TDWG codes: 79 MXE-DU MXN-SI MXS-JA MXS-NA
and refers to the fact that this species was previously
not recognized as distinct. Ecology
764
Vernacular names Pinus praetermissa occurs in open, dry pine-oak
woodlands or tropical broad-leaved forests, often
No vernacular names have been recorded for this on rocky slopes. Its altitudinal range is 9001900m
species. a.s.l. The dry season is from November to May,
annual precipitation varies between 10001500mm.
Description Associated pines are P. devoniana, P. lumholtzii, P.
pseudostrobus, and possibly P. oocarpa, but further
Trees to 1015(20)m tall, d.b.h. to 30cm; trunk investigation is necessary to evaluate its sympatry
monopodial, tortuous or curved, sometimes branch- with these and other pines.
ing low. Bark irregularly fissured longitudinally, red-
dish brown. Branches long, spreading, tortuous, not Conservation
pendulous, forming a broad, irregular, open crown.
Shoots slender, reddish brown, turning greyish. IUCN: NT
Cataphylls 710mm long, subulate, not recurved,
scarious, light brown. Vegetative buds oval-oblong Uses
to cylindrical; terminal bud 1015mm long; lateral
buds ovoid-acute, smaller, not resinous. Fascicle This scattered species is rarely recognized as distinct
sheaths 1014mm long, persistent, weathering dark from P. oocarpa by loggers and consequently no spe-
brown to grey. Leaves in fascicles of (4)5, persist- cific uses are known. It is likely to be exploited for
ing up to 3 years, straight, delicate and lax but not timber together with other species. It is a subtropical
pendulous, (8)1016cm long, 0.50.8mm wide, species not known in horticulture.
serrulate at margins, acute, bright green. Stomata on
all faces of the leaves. Pollen cones ovoid-oblong to
cylindrical, 11.5cm 5mm, pink to reddish. Seed Pinus pringlei Shaw, in Sargent, Trees & Shrubs
cones subterminal, commonly solitary, occasionally 1: 211, t. 100. 1905. Type: Mexico: Michoacan,
opposite, on up to 35mm long, recurved, slender Uruapn, Cerro de la Cruz, [hilltops near
peduncles, deciduous, non-serotinous, falling with- Uruapn], C. G. Pringle 10019 (lectotype GH).
out basal scales and peduncle in most cases. Mature
cones broadly ovoid to subglobose (wider than long), Etymology
often with a flattened base when opened and miss-
ing the proximal scales, (4)56.5(7) (5)68cm This species was named after Cyrus G. Pringle
when open. Seed scales thin woody but rigid, (18381911), who made many collections of plant
oblong, straight or recurved, more or less symmetri- specimens in Mexico.
cal. Apophysis flat to slightly raised, radially striate
or ridged to transversely keeled, rhombic to pen- Vernacular names
tagonal in outline, lustrous light brown or yellowish
brown. Umbo dorsal, flat or slightly raised, obtuse. Pringles pine; pino rojo (Spanish)
766
Plate 33. Pinus pseudostrobus. 1. Habit of tree. 2. Branchlets with foliage. 3. Leaves. 4. Seed cone (var.
pseudostrobus). 5. Seed cone (var. apulcensis).
recurved, reddish brown to dark brown. Vegetative habitat in cold-temperate to warm-temperate zones.
buds ovoid-conical, acute; terminal bud 1520 Its altitudinal range is considerable, but differs little
1015mm; lateral buds smaller, not resinous. along the entire 2500 km of its range: (850)1900
Fascicle sheaths (15)2030(35)mm long, persis- 3000(3250)m a.s.l. Annual precipitation is also
tent, lustrous red-brown, weathering grey-brown. variable, but with a minimum of ca. 800mm, in
Leaves in fascicles of 5, rarely 4 or 6, persisting 23 Guatemala and Honduras it can be over 2000mm.
years, slender, straight, spreading or drooping, usu- Throughout its range it is an important constituent
ally lax, (18)2030(35)cm long, 0.81.3mm wide, of mixed conifer pine and pine-oak forests, occa-
with serrulate margins, acute, (light) green or glau- sionally associated with Liquidambar, as on wet E
cous green. Stomata on all faces of leaves. Pollen slopes on the Gulf coast side of the mountains of
cones ovoid-oblong to cylindrical, 2035 57mm, Central and S Mexico. In the many disturbed forests 767
purplish- or pinkish yellow. Seed cones subtermi- it may survive as scattered groves or as individual
nal, solitary or in pairs, more rarely in whorls of trees, often with an understorey of e.g. Gaultheria,
34 on very short, stout peduncles remaining with Cassia or, when associated with fire, grasses and/
a few cone scales on branch when cone has fallen. or Pteridium aquilinum. In its driest habitat in
Mature cones more or less asymmetrical, often Central and NE Mexico it occurs with P. cembroides,
curved at base, ovoid-oblong to broadly ovoid when Juniperus flaccida, and Quercus and an understorey
opened, then 716 613cm. Seed scales usually with e.g. Agave, Buddleja, Opuntia, and Salvia, prob-
thick woody, oblong, straight or slightly curved. ably mostly in secondary forest.
Apophyses extremely variable, from nearly flat to
elongated, more so on one side of cone and towards Uses
base, transversely keeled, tapering to an obtuse umbo
(or umbo mucronate), rhombic or pentagonal in Pinus pseudostrobus is one of the most common
outline, upper margin angular, irregularly undulate, and important hard pines in the southern half of
or rounded; colour in various hues of brown. Umbo Mexico and the highlands of Guatemala and parts
dorsal, variable, from obtuse to prominent, 315mm of Honduras. Its wood is light yellowish in colour,
long, without a prickle or prickle deciduous. Seeds light weight (specific gravity 0.45), strong, with long
57 34.5mm, ochraceous to grey-brown, with intervals of knot-free wood which is only slightly
or without dark spots; wings obliquely-ovate, with resinous. Exploitation for timber is widespread
a straight side, 2025 710mm, yellowish brown. and has led to regional depletion of the best stands.
The wood is used for light construction, carpen-
Distribution try and joinery, wall panelling, veneers, boxes and
containers, matches, and wood pulp. In Mexico,
Mexico: Sinaloa-Durango border, Nuevo Len, SE this species is also used as a source of resin. This
Coahuila, E Guanajuato(?), Jalisco, Michoacn, tree has been widely planted for forestry in Africa
Mxico, Distrito Federal, Morelos, Hidalgo, Puebla, and to a lesser extent in India; in horticulture its is
Tlaxcala, W Central Veracruz, Guerrero, Oaxaca, rare and generally only suitable in regions with a
Chiapas; Guatemala (highlands); W Honduras; N El mild climate.
Salvador.
TDWG codes: 79 MXC-DF MXC-ME MXC-MO MXC- 2 varieties are recognized:
PU MXC-TL MXE-AG MXE-CO MXE-CU MXE-DU
MXE-GU MXE-HI MXE-NL MXE-QU MXE-SL MXE-
TA MXE-ZA MXG-VC MXN-SI MXN-SO MXS-CL MXS- Pinus pseudostrobus Lindl. var. pseudostrobus.
GR MXS-JA MXS-MI MXS-NA MXS-OA MXT-CI 80 Type: Mexico: Michoacan, Angangueo,
ELS GUA HON C. T. Hartweg s.n. (lectotype P).
Pinus pseudostrobus Lindl. var. apulcensis (Lindl.) Pinus oaxacana Mirov var. diversiformis Debreczy &
Martnez, Anales Inst. Biol. Univ. Nac. Mxico 16: Rcz, Phytologia 78 (3): 19. 1995.
192. 1945, non Shaw (1909); Pinus pseudostrobus
Lindl. subsp. apulcensis (Lindl.) Stead, J. Linn. Soc., Description
Bot. 89: 269. 1984, excl. typus.
Pinus nubicola J. P. Perry, J. Arnold Arbor. 68: 447. Apophysis variable, prominently raised and partly
1987. elongated, especially on one side of cone and towards
Pinus pseudostrobus Lindl. var. laubenfelsii Silba, base, transversely keeled, rhombic or pentagonal in
Phytologia 68: 60. 1990. outline; upper margin angular, irregularly undulate,
Pinus yecorensis Debreczy & Rcz, Phytologia 78 (3): or rounded; colour in various hues of brown. Umbo
768 15. 1995. dorsal, prominent and/or elongated, 515mm long,
Pinus yecorensis Debreczy & Rcz var. sinaloensis 510mm wide at base, without a prickle, usually
Debreczy & Rcz, Phytologia 78 (3): 18. 1995. darker than the apophysis.
Apophysis variable, from nearly flat to prominently For a taxonomic assessment and the correct applica-
raised, more so on one side of cone and towards tion of this infraspecific name and its synonyms, see
base, transversely keeled, tapering to an obtuse Farjon (1995) and Farjon & Styles (1997).
umbo, rhombic or pentagonal in outline; upper
margin angular, irregularly undulate, or rounded; Distribution
colour in various hues of brown. Umbo dorsal,
obtuse, 36mm long, 46mm wide at base, without Mexico: Mxico, Hidalgo, Puebla, Tlaxcala, W Central
a prickle or prickle deciduous, usually darker than Veracruz, Guerrero, Oaxaca and Chiapas; also in the
the apophysis. Guatemala highlands and in N El Salvador.
TDWG codes: 79 MXC-ME MXC-PU MXC-TL MXE-
Distribution HI MXG-VC MXS-GR MXS-OA MXT-CI 80 ELS GUA
Pinus oaxacana Mirov, Madroo 14: 145. 1958; Pinus Dwarf Siberian pine; Kiedrovnik (Russian)
pseudostrobus Lindl. var. oaxacana (Mirov) S. G.
Harrison, Taxon 14 (7): 247. 1965.
from fallen leaf fascicles, glabrous, yellowish green abandoned agricultural fields in many areas. In well
turning yellowish brown to darker reddish brown. established stands in the mountains it is often mixed
Buds ovoid to short cylindric, up to 10mm long, res- with Pinus rigida or P. virginiana (at lower altitudes)
inous, red-brown. Leaves in fascicles of 2, occasionally and the angiosperms Acer rubrum, Nyssa sylvatica,
3, held by a short, persistant basal sheath, spreading, Quercus prinus, Q. coccinea, Q. velutina, Castanea
persisting 3 years, (3)57(8)cm long, straight or dentata, and Oxydendrum arboreum. Several spe-
more or less contorted, rigid, 11.5mm wide, often cies of Rhododendron, Vaccinium, Gaylussacia, and
twisted, green; margins serrulate; apex acute to short Kalmia latifolia, Ilex montana and Smilax glauca
acuminate; stomata in fine lines on all surfaces. often form a dense understorey in stands of Pinus
Pollen cones in dense clusters, spirally arranged, pungens on steep mountain slopes.
770 ca, 1.5cm long, conspicuously red when growing,
turning yellow with the swelling of pollen sacs and Conservation
ripening of pollen. Seed cones lateral on branches,
rarely solitary, usually in whorls of 36, sometimes IUCN: LC
more, nearly sessile or on short peduncles, long per-
sistent, variously serotinous but eventually opening, Uses
asymmetrically ovoid-conical, 510cm long, 46cm
wide, opening widest near base to 8cm. Seed scales Table mountain pine is of low value as a timber tree
thick woody, cuneate from base; apophyses promi- due to small size and poor form, with crooked stems
nently raised, diamond-shaped at mid-cone, conical and long branches. Pulpwood and firewood are the
and recurved near base, transversely keeled, largest only uses commonly cited to have any commercial
on sun-exposed side of cone, orange-brown or yel- value. Its worth to society is more of an ecological
lowish brown; umbo dorsal and nearly central, stout kind, as protective forest cover on unstable, shallow
and armed with a dark, sharp spine. Seeds obovoid, rocky soils on mountain slopes and as habitat for
somewhat angular and flattened, 56mm long, wildlife. This species is not often seen in cultivation
blackish brown; wing 1525(30)mm long, brown. as it does not make a shapely tree; its gnarled form
on rocky outcrops in nature can be very picturesque
Distribution and reminiscent of Chinese or Japanese landscape
paintings. It is restricted to botanic gardens and
E USA: Appalachian Mts. and some outliers, mainly arboreta in Europe and North America.
in the Piedmont.
TDWG codes: 75 PEN WVA 78 GEO MRY NCA SCA
TEN VRG Pinus quadrifolia Parl. ex Sudw., U.S.D.A. Div.
Forest. Bull. 14: 17. 1897. Pinus cembroides Zucc.
Ecology var. quadrifolia (Parl. ex Sudw.) Silba, J. Int.
Conifer Preserv. Soc. 7 (1): 29. 2000. Type: USA:
Pinus pungens is found in the Appalachian Mountains California, San Diego Co., mountains E of San
from the foothills to the crests, it becomes the domi- Diego(Boundary Survey under direction of major
nant or single species of pine above 1300m a.s.l. in W. H. Emory), C. C. Parry 1390 (holotype US).
the southern part of its range. It grows on a variety Fig. 246
of soils, but is absent from limestone derived sub-
strates. In the surrounding broad-leaved forests it Pinus juarezensis Lanner, Southw. Naturalist 19 (1):
is restricted to outcrops or ridges of exposed rock. 77. 1974; Pinus cembroides Zucc. var. juarezensis
Precipitation levels vary from north to south and (Lanner) Silba, Phytologia 68: 48. 1990.
with altitude between 760mm and 2100mm annu-
ally; summers are cool to warm while winters are Etymology
moderately cold and wet. This pine is a pioneer spe-
cies, able to take advantage of sudden environmen- The species epithet refers to the common number of
tal changes, e.g. disturbance by forest fire; it invades four leaves in a fascicle.
Parry pinyon, Nut pine; pion (Spanish) The altitudinal range of Pinus quadrifolia is 900
2400(2700)m a.s.l. It grows between the semi-
Description desert and the chaparral scrub zones (and partly
within the latter) and the mixed coniferous forest on
Trees or large shrubs to 1015 m tall, d.b.h. to 3050 cm; the highest parts of the mountains. It is more widely
trunk short, erect, low branched or with a short, clear distributed and often more common than P. mono-
bole. Bark thick, rough and scaly, with deep lateral phylla in the Pinyon-Juniper woodland, but occurs
and longitudinal fissures, reddish brown; outer bark often with it. Pinus jeffreyi is the only other pine with
weathering grey. Branches spreading or ascending, which it occurs in Mexico. Juniperus californica and 771
the higher order branches slender, spreading or Quercus turbinella are common; in the chaparral zone
assurgent; crown rounded, becoming wide and open many shrubs, e.g. Adenostoma, Ceanothus, Artemisia,
in old trees. Shoots stout, rough with short decur- Cercocarpus, Rhus, Eriodictyon, Arctostaphylos, and
rent pulvini and small cataphylls. Vegetative buds Yucca, dominate. Most of these mountains are gra-
ovoid-conical; terminal bud 610 45mm; lateral nitic, but in the south of the ranges more volcanic
buds acute, orange-brown, slightly resinous. Fascicle rock is found. Pinus quadrifolia often grows in cracks
sheaths initially 58mm long, soon breaking up in among boulders. Annual precipitation is a moderate
recurving but early falling scales (not or rarely form- 300500mm, but it is very variable; most of it comes
ing rosettes). Leaves in fascicles of (3)4(5), rarely during winter cyclonic storms and there is a long dry
a few fascicles 2 or 6; fascicles persisting (3)47 season from spring through summer.
years; leaves curved or sometimes straight, rigid,
often connivent for some time, (1.5)24(5)cm Conservation
long, (0.8)11.5(1.7)mm wide, entire, acute-pun-
gent, dull or lustrous grey-green to glaucous green, IUCN: LC
with whitish stomatal bands. Stomata in two bands
on adaxial faces. Pollen cones ovoid-globose to short Uses
cylindrical, ca. 10mm long, purplish red, turn-
ing yellowish. Seed cones solitary or in whorls of There is no use of this species for timber. It is locally
24 on short, slender peduncles falling with cones. used for firewood. The seeds are edible and are har-
Mature cones ovoid-globose to globose when closed, vested to be sold in local markets. Resin may be
irregular when opened, 46 4.57cm when open. tapped on a small scale as well. It is not in cultiva-
Seed scales parting widely, moveable, irregular, con- tion outside a few botanic gardens and other collec-
cavo-convex, with 12 deep seed. Apophysis thick tions; it should be suitable in regions with hot, dry
woody, raised, pyramidal or obtuse conical, trans- summers and can endure frost and snow if not for
versely keeled, recurved or straight, ochraceous to prolonged time and of high altitude provenance.
yellowish brown or reddish brown, often resinous.
Umbo dorsal, flat or obtuse-pyramidal, centrally
indented. Seeds obliquely obovoid or elliptic, 1218 Pinus radiata D. Don, Trans. Linn. Soc. London 17:
812mm; integument thin (0.30.5mm); mega- 442. 1836.
gametophyte (endosperm) white. Seed wings rudi-
mentary on the scale, absent from the developed, Etymology
free seed.
The species epithet is said to refer to the radial mark-
Distribution ings on the apophyses of some cones.
Description Ecology
Small trees or shrubs to 39m tall, d.b.h. to 1540cm; The populations of Pinus remota are almost all
trunk short, contorted, soon branching. Bark thick, highly disjunct; the species has obviously retreated
rough and scaly, exfoliating with thin, scaly plates, to isolated mountains. Its altitudinal range is 1200
in old trees on lower part of trunk longitudinally 1600(1850)m a.s.l. (on the Edwards Plateau in
furrowed, grey to blackish grey. Branches spreading Texas it occurs considerably lower, the type collec-
or ascending, the ultimate branches stout, assurgent. tion is from 450m). It is restricted to canyons or
Shoots rough with prominent, non-decurrent pul- rocky mountain slopes, often on calcareous soil or
vini, which leave circular scars after fascicles have limestone rock, on dry sites where Pinyon-Juniper
774 fallen. Cataphylls ca. 5mm long, subulate, reflexed, woodland is not well developed and the pines often
grey. Vegetative buds ovoid to ovoid-cylindric; ter- remain shrubby. Annual precipitation ranges from
minal bud 57mm long; lateral buds shorter, acute, 300500mm, but is extremely variable from year
not resinous or sometimes slightly resinous. Fascicle to year. Frost is common in December and January.
sheaths initially ca. 5mm long, soon caducous, not This species occasionally occurs with P. cembroides
recoiling to form a rosette. Leaves in fascicles of and more rarely with P. arizonica var. stormiae, com-
2(3), persisting 45 years, slightly curved to falcate, mon are Juniperus monosperma and J. ashei (in the
rigid, (2)34.5(5.5)cm long, 0.81.1mm wide; northern part of the range), Quercus, Cercocarpus
margins entire; apex acute-acuminate; abaxial face and semi-desert plants e.g. Agave lecheguilla,
dull light green or yellowish green; adaxial face(s) Opuntia and Fouquieria splendens.
slightly glaucous, with inconspicuous whitish sto-
matal lines. Stomata on all faces of the leaves. Pollen Conservation
cones subglobose to ovoid, 45mm long, pink or
purplish, turning light yellow. Seed cones solitary or IUCN: LC
sometimes in pairs on 58mm long, slender, curved
peduncles, maturing in two years. Mature cones Uses
subglobose or globose when closed, (2)2.54
36cm when open. Seed scales parting widely; basal Any use of this species (firewood, edible seeds)
scales remaining connate; all scales loosely attached is probably incidental or at most on a very small
to the rachis, irregularly shaped, concavo-convex, scale due to its scattered occurrence in often
with incurved margins and 12 deep seed cavities. remote places. The edible seeds are harvested and
Apophysis prominently raised, transversely or radi- sold with those of other pinyons. It is not known
ally keeled, irregularly rhombic to pentagonal in to be in cultivation outside a few botanic gardens,
outline; upper margin angular and irregular; colour but should be well adapted to gardens with a dry,
(lustrous) light brown to reddish brown. Umbo hot summer.
dorsal, flat or indented, with a minute, deciduous
prickle. Seeds obovoid-angular or ellipsoidal, 1216
810mm, light ochraceous with a grey tinge; integ- Pinus resinosa Aiton, Hort. Kew. 3: 367. 1789. Type
ument very thin, 0.10.4mm; megagametophyte not designated.
(endosperm) white. Seed wings vestigial, remnants
remain on the scale when the seeds are dispersed. Etymology
Description Ecology
Trees to 37m tall; trunk to 1.5m d.b.h., straight, Pinus resinosa is a species of pine occupying the
shorter and with heavy branches in solitary trees. northeastern mixed conifer-deciduous broad-leaved
Bark on young trees and in crown of older trees thin, forest bordering on the boreal conifer forest at its
orange-brown, on trunk becoming thick and break- northern limit. This area was heavily glaciated dur-
ing into irregularly shaped but wide, elongated, ing the last Ice Age and the glaciers left huge depos-
scaly, reddish brown plates divided by grey-brown its of sand, some of which in post glacial time was
fissures. Branches spreading wide, forming a blown into sand dunes. It is to these sands that this
rounded crown. Foliage branches moderately slen- pine is largely confined, as here it can successfully
der, new shoots to 1cm thick, light orange-brown compete with other trees. Another type of habi- 775
or red-brown, turning darker brown, rough with tat for P. resinosa is swamp margins. The species
pulvini from fallen leaf fascicles. Buds ovoid to avoids non-acidic soils and only grows over lime-
cylindrical; terminal bud to 20mm long, resinous; stone when there is a leached, acid top layer. It is
cataphylls red-brown, fringed, some free at the apex. largely a lowland pine growing at altitudes between
Leaves in fascicles of 2, held in a basal sheath ini- 200m and 450m, but ascends to 1300m a.s.l. in
tially 1520mm long, but eroding to basal part of the Appalachian Mountains. In the northern parts
only 56mm length, persisting 34 years, straight of its range it can form extensive pure stands, but
or curved, 1218cm long, slightly twisted, brittle, more often it is growing together with Pinus banksi-
breaking easily, ca. 1.2mm wide, dark green; mar- ana, P. strobus, or both. Populus spp. are common in
gins minutely serrulate; apex abruptly acute; stomata the sub-boreal zone, further to the south Abies bal-
in fine lines on all surfaces. Pollen cones clustered at samifera, Acer rubrum, A. saccharum, Quercus alba,
base of new shoots, spirally arranged, ovoid-oblong, Q. rubra, Fagus grandifolia, and Betula papyrifera
1.5cm long, conspicuously red or purple to dark can come into the forest, especially on more fertile
purple, turning dark brown shortly before they fall. soils. In mountainous areas, Pinus rigida, P. pungens,
Seed cones solitary or in pairs, nearly sessile, fall- P. virginiana, and Tsuga canadensis are other conifers
ing without the short peduncle and often leaving growing with P. resinosa. Where this pine is attaining
a few basal scales behind, 46cm long, narrowly good size most of the associated tree species remain
ovoid-conical when closed, symmetric or slightly lower, but in some of the broad-leaved forest types
asymmetric at base, opening soon after matura- P. resinosa is only a minor component restricted to
tion to a broadly ovoid or sub-globose shape with the poorest sites. This pine is dependent on fire for
a more or less level base. Seed scales thin woody, its natural reproduction and is therefore a seral spe-
rigid, oblong, dark brown; apophyses slightly raised, cies on most sites.
transversely keeled, sometimes more pronounced
on exposed side of cone near base; umbo central, flat Conservation
or depressed, unarmed or with a minute, deciduous
prickle. Seeds ovoid, 45mm long, slightly flattened, IUCN: LC
rugose, brown; wing 1520mm long, narrow, pale
brown. Uses
This species occupies the cool maritime and partly Pinus roxburghii Sarg., Silva N. Amer. 11: 9. 1897.
mountainous NE of the USA, where it occurs from Type: India: Himalaya, [locality not given],
sea level in the north to nearly 1400m in the south- W. Roxburgh B-W 17762 (holotype not located,
ern Appalachians. It grows on shallow sandy or isotype B-W).
gravelly soils poor in nutrients, usually well drained,
but sometimes water-logged, as in the swamps of Etymology
New Jersey. It is a seral species, most commonly
associated with oaks (Quercus spp.) or with other The species name commemorates William Roxburgh
pines e.g. P. virginiana, P. echinata, or P. pungens. (17511815), a Scottish botanist working for the East
In swamp areas it grows with Chamaecyparis thy- India Company and Superindentent of the Calcutta
oides. On some sandy sea shores (e.g. at Cape Cod, Botanic Garden.
Massachusetts) a decumbent, wind-swept form
15m tall occurs. It is exceptional in its capacity to Vernacular names
resprout from stumps; small tufts of adventitious
foliage appear on the trunks even of healthy trees. Its Chir pine, Long-leaved Indian pine; Chil (Hindi);
ability to regrow foliage after severe damage at any Dhup (Nepalese); Chir, Sula (Pakistan)
stage in its development enables this species to sur-
vive fires as well as browsing of seedlings and sap- Description
lings. In areas with regular occurrence of fire many
trees are multi-stemmed from a basal stool. Trees to 5055m tall, usually not taller than 30m;
trunk to 3m d.b.h., usually to 1m d.b.h., bole
Conservation straight and columnar. Bark becoming thick and
divided by long, deep fissures, breaking into large,
IUCN: LC elongated, scaly plates, weathering grey-brown, with
dark brown and purplish hues. Branches self-prun- est growing, scattered individuals at 2500m. Pinus
ing, long and spreading to ascending, forming an roxburghii is restricted to the monsoon belt with
open, domed crown in mature to old trees. Foliage summer rains. In its higher altitudinal range this
branches slender or stout, pale grey or light brown, pine species is commonly mixed with Cedrus deo-
covered with leaf-like brown cataphylls remaining dara and Pinus wallichiana, but occurs below the
several years and becoming recurved at their apices. forest zone characterized by species of Abies. Broad-
Buds small, ovoid, without resin. Leaves near ends of leaved trees (angiosperms) are commonly Quercus
branchlets, persisting 12 years, in fascicles of 3, held incana, Schima wallichii and Rhododendron arbo-
together by persistent, 2530mm long basal sheaths, reum. Towards its lower limit angiosperms become
spreading and slightly drooping, 2530(35)cm long, more dominant and the pines occur on rocky slopes
778 pliant, slender, broadly triangular in cross-section, with a northern or eastern aspect.
1.21.7mm wide; margins minutely serrulate; leaf
colour bright green; apex thinly acute-acuminate; Conservation
stomata in fine lines on all surfaces. Pollen cones
clustered near base of new shoots, spirally arranged, While forest destruction and logging have reduced
ovoid-oblong, 1315mm long. Seed cones solitary or the area of occupancy (AOO) of P. roxburghii, it is
in whorls of 25 on stout branches, short peduncu- still covering extensive areas (an estimated 0.87mil-
late, persistent, heavy, broadly ovoid or ovoid-coni- lion ha in India alone) and is therefore not consid-
cal when olive green, growing to 1015(20)cm long, ered to be threatened with extinction.
712cm wide when closed, opening only slightly IUCN: LC
after several years to 13cm max. width. Seed scales
oblong, woody and rigid; apopyses strongly devel- Uses
oped, thick, conical, with rhombic to irregularly pen-
tagonal bases, sharply transversely keeled, (strongly) Chir pine is an important pine for resin production
recurved, smooth, lustrous yellowish brown or grey- in the Himalayan region, especially in NW India. It
brown; umbo triangular, obtuse and unarmed. Seeds has been the basis of the Indian naval stores indus-
obovoid, 812(15)mm long, slightly flattened; wing try, initiated by the Indian Forest Department under
adnate, oblique, 2025mm long, 810mm wide, British rule in 1888 (Langenheim, 2003) to sup-
lighter brown than the seed. ply the British Empire with turpentine and related
products. Later, it became the main source of these
Distribution substances for India, but production was falling dra-
matically due to poor management of the forests
Himalaya, from Pakistan to NE India, Arunachal and destructive tapping methods. In recent years
Pradesh (Assam, Kameng District). these have improved under more rigorous manage-
TDWG codes: 36 CHT 40 EHM-AP EHM-BH EHM-DJ ment. Artificial camphor is the main end product
EHM-SI NEP PAK WHM-HP WHM-JK WHM-UT derived from resin of P. roxburgii in the region; it is
also used for medicinal treatments. The wood is of
Ecology importance for railway sleepers after treatment for
preservation, and for construction, carpentry and
Pinus roxburghii is widespread and common in the joinery; it is also pulped for the paper industry. The
north-south oriented outer valleys of the Himalaya bark has a high tannin content (1114%) and is used
and its foothills and often forms pure stands espe- for tanning leather and for staining wood to give it
cially on dry, fire-prone slopes. Mature trees are an orange colour. The seeds are edible but not very
relatively fire resistant; regeneration after destruc- good. The needles are used for animal bedding and
tive fires can be massive and rapid when it acts as mixed with manure serve as a traditional fertilizer
a pioneer species. In prolonged dry seasons it may for agriculture. This species is uncommon in cultiva-
drop most of its leaves. It occurs on a variety of sub- tion outside India and Pakistan, but has been intro-
strates, from deep soil to bare rocks. Its altitudinal duced to South Africa as a forestry plantation tree. It
range is from 400m to 2300m a.s.l., with the high- is sometimes seen as an ornamental tree in countries
around the Mediterranean Sea. Despite the fact that closed, ovoid when opened, usually symmetrical,
in its natural habitat P. roxburghii does not occur in slightly flattened at base, 1015 68.5cm when
the Himalayan frost zone, some provenances could open, very resinous. Seed scales thin woody, rigid,
prove to be hardy. oblong, concavo-convex, often wider than apophy-
sis. Apophysis prominent, similarly shaped around
cone, transversely keeled, rhombic or pentagonal in
Pinus rzedowskii Madrigal & M. Caball., Bol. Tcn. outline, turning ochraceous or light brown. Umbo
Secr. Agric. Ganad. Subsecr. Forest. Fauna 26: 1. dorsal, rhombic-pyramidal, transversely keeled,
1969. Type: Mexico: Michoacan, Coalcomn de obtuse or with a minute prickle. Seeds (6)8(10)
Matamoros, Dos Aguas Forest Station, Pinabete, X. (4)56mm, brown; wings 2030(35) 813mm,
Madrigal 2202 (holotype INIF). Fig. 247, 248 brown, with darker stripes. 779
Etymology Distribution
This species was named after the botanist Jerzy Mexico, W Michoacan (municipality Coacomn,
Rzedowski, a student of the Mexican flora. W and SW of Dos Aguas in 1012 localities).
TDWG codes: 79 MXS-MI
Vernacular names
Ecology
Rzedowskis pine
The two smaller classical subpopulations, Cerro de
Description Chiqueritas and Cerro Ocotoso, are on steep talus
of large, eroded limestone blocks, near the summits
Trees to 1530m, d.b.h. to 3060cm; trunk mono- of small mountains in the mainly volcanic Sierra
podial, erect, often curved or contorted. Bark up to Madre del Sur. Each have only from 10 to a few score
56cm thick on lower part of larger trunks, rough trees, from old to saplings. Some subpopulations
and scaly, breaking into large, exfoliating plates are on more level ground, also with limestone boul-
divided by deep, longitudinal fissures, dark brown, ders, but interspaced with other rocky substrates.
weathering grey. Branches spreading and ascend- In the two former areas, the trees remain small, less
ing, often contorted; crown pyramidal in young than 15m tall; in some of the larger subpopulations
trees, open and irregular in old trees. Shoots slender, trees to 30m have been found. The altitudinal range
glabrous and smooth, greyish. Cataphylls ca. 5mm is 17142480m a.s.l. Annual precipitation is ca.
long, fragile, soon deciduous. Vegetative buds ovoid- 1500mm, most of it occurs from June to October.
oblong; terminal bud 810 45mm; lateral buds The climate is warm-temperate, with a minimum of
of nearly equal size, not resinous. Fascicle sheaths 5 C (December) and a maximum of 30 C (April).
of young leaves 79mm long, soon recoiling, form- Although surrounded by extensive mixed pine for-
ing a rosette at base of fascicle, finally deciduous est with species like P. pseudostrobus, P. herrerae and
before fascicles. Leaves in fascicles of (3)45, per- P. oocarpa, these species do not grow on the lime-
sisting 23 years on branchlets, straight or slightly stone talus. There, Quercus and shrubs, e.g. Clusia
reflexed, lax but not drooping, 610cm long, 0.6 salvinii, form an understorey with Agave and tall
0.8mm wide, with irregularly serrulate margins, herbs. Fires occur frequently, but rejuvenation
acute-acuminate, yellowish green or greyish green, seemed good at least at Cerro Chiquerita visited by
weakly glaucous on adaxial faces. Stomata on the me in 1994.
two adaxial faces only. Pollen cones after shoot elon-
gation forming a 57cm long spike leaving shoot Conservation
base and apex free; cones small, 5 3mm, purplish,
turning brown. Seed cones solitary or in whorls of The population is small and consists of 12 subpop-
24 on 1530mm long peduncles which fall with ulations, ranging from 1 to 3500 individuals, total-
cones. Mature cones ovoid to ovoid-conical when ing 60006500 individuals (Delgado et al., 1999).
Regeneration is quite abundant at some of the sites. a broad, irregular and open crown. Shoots slender
From this it is here inferred that the total of mature or stout, rough with prominent, decurrent pulvini
trees is probably around 1000. There is consider- from fallen leaf fascicles, pale grey-brown becom-
able genetic diversity, indicating a larger population ing darker brown. Buds ovoid-conical; terminal bud
in the (geological?) past (Delgado et al., 1999). The 1525mm long; lateral buds smaller, all resinous,
greatest threat to this species would appear to be a acute; cataphylls appressed, reddish brown, with
forest fire, as it is not a fire-adapted pine. Its occur- paler fringed margins. Leaves in fascicles of 3, held
rence amidst extensive pine forests make it vulner- by persistent, initially 20mm long but later much
able to this hazard. There is evidence in several of reduced (57mm) sheaths, spreading or drooping,
the scattered stands of ground fires having occurred persisting 34 years, (15)2028(32)cm long, pliant,
780 (Perry, 1991). slightly twisted, 1.5mm wide; margins serrulate; apex
IUCN: VU (D1, 2) acute-pungent to subulate; leaf colour grey-green;
stomata in conspicuous lines on all faces of leaves.
Uses Pollen cones ovoid to ellipsoid, 1015mm long, yel-
low becoming orange-brown. Seed cones at base of
No uses are recorded of this species. It is a botanical a subsequent leading shoot, solitary or occasionally
rarity only present in a few botanic gardens and pri- in pairs, on short, persistent peduncles, holding the
vate collections. Its altitudinal range implies occa- cone to the branch for up to 7 years and retaining a
sional light frosts, so it might prove hardy in mild few basal scales when falling; mature cones broadly
regions and as an attractive tree should be worthy of ovoid, massive, nearly symmetrical, opening slowly,
cultivation, which would also help its conservation 1725 1520cm when open, more or less flat or
as a species. convex at base, extremely resinous. Seed scales thick
woody and rigid, widest towards the apophysis,
with two seed cavities on adaxial side, dull brown;
Pinus sabiniana Douglas ex D. Don, in Lambert, apophyses very strongly developed, thick woody,
Descr. Pinus, ed. 8, 2: p. s.n. inter 144 et 145, t. 80. sharply transversely keeled, more or less abruptly (or
1832. Type: USA: California, D. Douglas s.n. 1826 gradually)merging into a long, curved umbo, up to
(holotype not located, isotypes GH, K [ex Herb. 20mm wide, chocolate-brown. Umbos dorsal, elon-
Hort. Soc. London]). gate, curved, with keeled sides, 1020mm long, to
12mm wide at base, ending in a sharp uncinate claw.
Etymology Seeds narrowly obovoid, slightly flattened, 1520
710mm, smooth, dark brown; wings short and
This species was named after Sir Edward Sabine, for- wide, ca. 10mm long, largely ineffective.
mer President of the Royal Society.
Distribution
Vernacular names
USA: California (lower slopes and mountains
Gray pine, Bull pine, Digger pine (deprecated) around Central Valley), Oregon (Jackson Co.).
TDWG codes: 73 ORE 76 CAL
Description
Ecology
Trees to 25m tall, usually remaining smaller; trunk
to 1m d.b.h., monopodial or forked, straight or Pinus sabiniana grows in the summer-dry moun-
curved. Bark on trunk thick, rough, scaly, divided tains and foothills that encircle the Central Valley of
into irregular plates and deep, longitudinal fissures, California, from the edge of the Mojave Desert to the
brown-grey or blackish grey with reddish brown fis- slopes above the Pacific Ocean. Its altitudinal range is
sures. Branches of first order long, spreading hori- from 50m to 1800m a.s.l. Annual precipitation var-
zontally or ascending, usually persistent along much ies much within its range, from 250mm per annum
of trunk; branches of higher orders sparse, forming near the desert to 1780mm at its upper limits in the
Sierra Nevada. It grows inland from the coastal fog Vernacular names
belt and does not tolerate hard frosts. Near the coast,
it grows in the chaparral zone with ericaceous shrubs Pond pine, Marsh pine, Pocosin pine
which is subject to frequent fires. On the lower slopes
of the Sierra Nevada it grows in the Upper Sonoran Description
Life Zone, mainly accompanied by various species of
Quercus, as it does at higher elevations in the Coast Trees to 20m tall; trunk to 60cm d.b.h., straight or
Ranges. Here Pinus coulteri can be an associate, in often crooked. Bark thick and breaking into irreg-
the north of its range it grows with Juniperus occi- ularly shaped, flat but scaly, dark reddish brown
dentalis. The woodlands with P. sabiniana are usually plates divided by deep fissures. Adventitious foliage
very open, with trees emering from shrubs or stand- shoots common in crown on largest branches, few 781
ing in areas covered with sparse grasses and herbs. on trunk; primary branches spreading and ascend-
The heavy cones are predated by squirrels and jays, ing, tortuous, poorly self pruning, forming an irreg-
the former can detach cones from the trees and gnaw ular and thin crown. Foliage branches sparse, stout,
through the thick scales to obtain the seeds, the latter new shoots 1cm thick, yellowish orange, frequently
play a role in seed dispersal and germination. glaucous, turning brown, forming numerous buds.
Buds narrowly ovoid to cylindrical; terminal bud
Conservation 1520 mm long, extremely resinous; cataphylls
appressed, red-brown, fringed; apex cuspidate.
IUCN: LC Leaves in fascicles of 3, occasionally 4 (adventitious
shoots can have dwarf shoots with 35 leaves), held
Uses in a basal sheath initially 2025mm long, reducing
to ca. 10mm, persisting 23 years in tufts at the ends
Gray pine nowadays has little value as a timber tree; it of branchlets, straight, (13)1520(21)cm long,
only played a major part during the California Gold flexible, slightly twisted, 1.31.5(2)mm wide, green;
Rush period due to its proximity to the gold fields. margins minutely serrulate; apex acuminate; stomata
Native American tribes used the seeds for food and in fine lines on all surfaces. Pollen cones clustered
the copious resin to seal drums and baskets. The at base of new shoots, spirally arranged, cylindric,
nutritional value of the seeds can well compete with 23cm long, yellowish brown. Seed cones solitary or
most other edible pine seeds, yet there is little or no in whorls of 25, sessile or on a 1cm long peduncle,
commercial use at present. The wood, because of long persistent, variously serotinous or opening in
its irregular size and shape and high resin content, the second year, (5)610cm long, ovoid-conical
is only used for railroad sleepers, pallets and wood when closed, symmetric, opening to a broadly ovoid
chips. The needles and twigs yield oils and turpen- or subglobose shape to 8cm wide with a flattened
tine. Its irregular, open crown and twisted growth, base. Seed scales thin woody, rigid, oblong, dull
usually making several stems, discourages use as an brown with a dark red-brown sealing band; apophy-
amenity plantation tree; its only interest is in arbo- ses slightly raised, more or less rhombic, transversely
reta and pineta where the climate is suitable, e.g. in keeled, light brown or pale red-brown; umbo dor-
the south of England, western France, around the sal and central, small, armed with a short, weak
Mediterranean, or in Australia. prickle, sometimes unarmed. Seeds obliquely ellip-
soid, 56mm long, slightly flattened, pale brown, or
mottled dark brown; wing 1520mm long.
Pinus serotina Michx., Fl. Bor. Amer. 2: 205. 1803.
Type not designated. Taxonomic notes
character of adventitious shoots. There are sufficient thirds of its height. Its wood is therefore mainly used
independent character differences to separate the as pulp wood. Slow growth in the nutrient poor
two species, although several of these are quantita- acidic swamps makes commercial use unlikely; how-
tive (to do with measurements) rather than qualita- ever, its growth is considerably better in plantations
tive (absent or present). Distribution and ecology of on well drained sandy soils. It is of limited value
the two species only overlap marginally. in amenity planting and horticulture and attempts
at forestry plantations in China, South Africa and
Distribution Zimbabwe have generally had poor results com-
pared to other pine species.
SE USA: from New Jersey to Alabama and Florida
782 (coastal plain).
TDWG codes: 75 DEL MRY NWJ 78 ALA FLA GEO Pinus sibirica Du Tour, in Dterville, Nouv. Dict.
NCA SCA VRG Hist. Nat. 18: 18. 1803. Pinus cembra L. var. sibirica
(Du Tour) G. Don, in Loudon, Hort. Brit. 1: 387.
Ecology 1830. Pinus cembra L. subsp. sibirica (Du Tour)
Krylov, Fl. Altai. Tomsk. 7: 1724. 1914 [& in Fl. Zap.
Pinus serotina is a lowland pine of wet, swampy areas Sibir. 1: 77. 1927]. Type not designated.
on the coastal plains of the Atlantic Ocean and the
Gulf of Mexico. It occurs on sandy flats with a high Pinus hingganensis H. J. Zhang, Bull. Bot. Res.
water table and in swamps. The climate on these North-East. Forest. Inst. 5 (1): 151. 1985; Pinus sibirica
plains is mild and humid with annual precipitation Du Tour var. hingganensis (H. J. Zhang) Silba,
ranging from 1120mm to 1420mm, with the dri- Phytologia 68: 61. 1990.
est months in winter. In so-called pocosins, peat
swamps that have risen above the water table and Etymology
drain outwards, P. serotina is common (a local ver-
nacular name is Pocosin pine). Ponds are swamps The species epithet refers to its natural occurrence
in depressions with very poor drainage. In both in Siberia.
types of swamp the pines form a tall, open canopy,
under which is a dense and varied shrub layer char- Vernacular names
acterized by Smilax laurifolia. Pinus serotina is often
associated with Taxodium distichum or with other Siberian pine, Siberian stone pine; Sibirskii kedr,
pines, e.g. P. taeda and P. elliottii. Common broad- Kedr (Russian)
leaved trees (angiosperms) belong to the genera
Nyssa, Magnolia, Liriodendron, Persea, and Ilex. Its Description
capability to resprout differs from that of P. rigida in
that adventitious shoots appear more in the crown Trees to 3540m tall; trunk to 1.8m d.b.h. Bark
than on the trunk, probably because fires tend to be smooth and pale brown on young trees and on
crown fires in the wetter forests with water-logged branches, becoming scaly and deeply fissured on
forest floors where this species occurs. trunks of large trees, turning grey. Branches numer-
ous, spreading and assurging, forming a dense,
Conservation broadly conical crown. Foliage branches slender to
stout, initially with a dense, pale yellow pubescence,
IUCN: LC then glabrous, smooth, yellowish brown or light
brown. Buds ovoid-conical, without resin; cataphylls
Uses reddish brown. Leaves in dense tufts towards end of
shoots, persiting 24 years, in fascicles of 5 held by
The wood of Pond pine is coarse grained and resin- deciduous basal sheaths of thin, orange-brown scales
ous, while the tree only attains modest size and is that fall away in the second year, spreading wide or
often crooked or divided into long branches at two forward, straight or slightly curved, more or less rigid
to flexible but not lax, 611(13)cm long, triangular as well as birds and other animals; the Eurasian
in cross-section, not twisted, 1.21.7mm wide; mar- Nutcracker (Nucifraga cariocatactes, fam. Corvidae)
gins minutely serrulate; leaf colour green; stomata is almost entirely responsible for the effective disper-
in white lines on the two adaxial faces. Pollen cones sal of the seeds by making food caches.
clustered, spirally arranged, short cylindrical, initially
reddish turning red-brown. Seed cones intitially Conservation
erect, becoming patent, single or in 23 in whorls on
very short peduncles, remaining closed or opening IUCN: LC
only slightly at maturity, from glaucous green to pur-
plish when growing, resinous, ovoid-conical, usu- Uses
ally longer than wide but variable, (5)710(12)cm 783
long, 46(8)cm wide, dark brown when ripe. Seed The wood of Siberian pine is considered inferior in
scales imbricate, widely cuneate proximally, with 2 quality to that of Scots pine and Siberian spruce,
deep seed cavities adaxially, soft woody; apophyses two of the (co)dominant conifers in Russia/Siberia.
broadly rhombic or widely triangular to semi-orbic- However, it still serves numerous uses, such as round-
ular, thickened, pilose when growing, longitudinally wood for poles, sawn timber for light construction,
striated when full grown and drying to dark brown; carpentry, furniture, veneers, utensils, boxes, wood
umbo terminal, obtuse, slightly upturned, lighter carving, and musical instruments. The wood is soft,
coloured than the rest of the scale. Seeds oblong- lightweight and rose-coloured with a good texture
obovoid, 1014 57mm, slightly ridged across one for finer applications. Trees are also tapped for resin,
end but without a true wing. mainly to produce turpentine. The edible seeds have
a very high fat content (ca. 65%) and contain many
Distribution vitamins. In the Altai Mountains harvests may yield
200300 kg of nuts or kernels (= seeds) per ha.
China: Heilongjiang (Tuqiang), Nei Mongol, Xin The seeds have to be separated from the cones with a
jiang; Kazakhstan; Mongolia; Siberia; an isolated mechanical cone-thresher as the cone scales do not
population is reported from the Kola Peninsula in open to release the seeds (which in nature is done
NW Russia. by the Eurasian Nutcrackers strong bill). In Russia
TDWG codes: 14 RUE RUN 30 ALT BRY CTA IRK KRA this species is commonly planted as an amenity tree
TVA WSB YAK 32 KAZ 36 CHI-NM CHM-HJ CHX 37 in large gardens and parks. Few cultivars are known,
MON but in Russia several dwarfed growth forms found
at high altitude or in peat bogs have been described
Ecology as botanical forms and may also be in cultivation. In
Europe, the closely related Arolla pine (P. cembra) is
This is a species growing both in lowland areas along more commonly planted.
the great Siberian rivers like the Ob and the Yenisei
Rivers between 100m and 200m a.s.l., and in the
mountains to an altitude of around 2400m, where it Pinus squamata X. W. Li, Acta Bot. Yunnanica 14
forms dense, pure forests. In the lowlands, it occurs (3): 259. 1992. Type: China: Yunnan, Qiaojia Xian,
with Pinus sylvestris, Larix gmelinii or L. sibirica and X. W. Li 91250 (holotype SWFC).
Betula pendula on drier sites that regularly burn and
with Abies sibirica and Picea obovata (dark conifer- Etymology
ous forests) and Betula sp. on more mesic sites in
the river basins, in bog margins, and uplands. Pinus The species epithet (Latin squamata = furnished
sibirica is one of the major forest-forming conifers with scales) refers to the scaly bark of larger trees.
in the Siberian taiga and is estimated to cover ca.
45million hectares. It is the slowest growing spe- Vernacular names
cies in these extensive forests and may live up to 850
years. Its edible seeds are harvested by local people qiao jia wu zhen song (Chinese)
Pinus strobiformis Engelm., in Wislizenus, Mem. cylindrical to ovoid-oblong when opened, 1230(
Tour N. Mexico: 102. 1848. Pinus ayacahuite 60) 711cm when open. Seed scales spreading
Ehrenb. ex Schltdl. subsp. strobiformis (Engelm.) obliquely to patent; basal scales recurved or reflexed;
E. Murray, Kalmia 13: 21. 1983. Type: Mexico: most scales with curved margins, thick woody, of
Chihuahua, Cosiquiriachi, F. A. Wislizenus 231 similar shape around the cone but differentiating
(holotype MO). from base to apex, with one or two deep seed cavities
adaxially. Apophysis more or less cuneate or elon-
Pinus strobiformis Engelm. var. carvajalii Silba, gated, thick at the proximal end, thinning out and
Phytologia 68: 61. 1990. straight, recurved or reflexed distally, smooth or lon-
Pinus ayacahuite Ehrenb. ex Schltdl. var. novogali- gitudinally furrowed, often very resinous, light yel-
ciana Carvajal, in McVaugh, Fl. Novo-Galiciana 17: lowish brown or dark ochraceous. Umbo terminal, 785
48. 1992. broadly triangular, obtuse. Seeds obovoid, 1218
811mm, reddish brown or brown. Seed wings vesti-
Etymology gial to very small, up to half the length of the seed in
some cones.
The species epithet means shaped like strobus, i.e. it
is compared with Pinus strobus. Distribution
2000m a.s.l. Annual precipitation varies greatly Pinus strobus L. var. strobus. Type: North America:
from area to area, with lows at around 500mm [locality not stated] 5 K Strobus, P. Kalm LINN
and highs in Mexico to 3000mm. The southern 1135.10 (lectotype LINN).
var. chiapensis experiences no frost, while long and
cold winters are the norm in most of the range of Description
var. strobus. Both varieties are major or minor com-
ponents of mixed forests, with other conifers and/ Seed cones (8)1018(20)cm long.
or with broad-leaved trees. There is a similarity of
several broad-leaved (angiosperm) tree species in Distribution
the forests of the southern Appalachians and the
mountains of Veracruz and Chiapas, Mexico, but E North America: from Newfoundland to N Georgia, 787
in the colder north P. strobus grows with species westward to Minnesota.
not common to both the northern and southern TDWG codes: 72 74 75 78
ranges.
Conservation
Conservation
The vast resources of timber available to European
See under varieties colonists from this large pine had been depleted
towards the end of the 19th century. However, as
Uses regrowth occurred, this has not threatened the con-
tinued existence and occurrence of the species signif-
Eastern white pine was once the most important icantly. Hence, while old growth Eastern white pine
timber tree in eastern North America and in colonial is now very rare, under IUCN criteria this variety of
times the British government forbade European col- P. strobus is not under threat.
onists to cut the larger trees (marked with the broad IUCN: LC
arrow) as it wished to reserve these for the British
Navy as ship masts. Its fine grained, smooth textured
wood low in resin makes excellent construction Pinus strobus L. var. chiapensis Martnez, Anales
timbers, while doors and windows, furniture, and Inst. Biol. Univ. Nac. Mxico 11: 81. 1940. Pinus
matches are other uses. In the USA and Canada it is chiapensis (Martnez) Andresen, Phytologia 10: 417.
widely planted both for timber and for urban plant- 1964; Pinus strobus L. subsp. chiapensis (Martnez)
ing as shelter belts, as well as restoration of areas E. Murray, Kalmia 12: 23. 1982. Type: Mexico:
disturbed by strip mining of coal. Americans and Chiapas, Ocotepec, M. Martnez s.n. (lectotype
Canadians use this pine for Christmas trees as its MEXU).
foliage can be clipped into shape. As an ornamental
tree it is not very common in Europe, probably due to Description
susceptability to White pine blister rust (Cronartium
ribicola, Basidiomycota) and poor shape in cultiva- Seed cones highly variable in length, 625cm.
tion. Attempts at forestry plantation in Britain also
failed due to this disease, as well as to aphid insect Taxonomic notes
predation (Pineus strobus). In the colder, drier win-
ters of North America it thrives better; several culti- While Mrtinez (op. cit.) described and named this
vars are well known and used in gardens, especially taxon as a variety of P. strobus as it has only insig-
some of the dwarfed forms. nificant morphological character differences with
that species, some others have treated it as a spe-
2 varieties are recognized: cies. It does not have smaller cones as these are
very variable in size and no discontinuous charac-
ter states have been found between this variety and
var. strobus. Statistical analysis applied to continuous
character states to demonstrate significant differ- ing grey, flaking in small or large chips. Branches
ences between the two taxa (Andresen, 1966) is in my spreading or curving down to pendulous, contorted,
view an unconvincing approach to the species prob- forming domed or irregular, open crowns in most
lem. Its considerable distance (2400 km) from var. cases. Foliage branches slender, spreading or pendu-
strobus, which can be explained by late Pleistocene lous in lower crown, becoming rough with pulvini
climatic events, is not a sufficient reason to assign after shedding leaves, sometimes remaining smooth,
it the rank of species. If the disjunction persists, it glabrous, new shoots yellowish green, grey-green or
may evolve truly distinct characters and become a pale yellow, turning light brown to grey-brown or
full species eventually. grey. Buds ovoid or ellipsoid, acute; terminal bud to
15mm long; lateral buds smaller, all without resin
788 Distribution or slightly resinous; cataphylls red-brown or pale
brown. Leaves in fascicles of 2, more or less remote,
Mexico: in Guerrero, E Puebla, Veracruz, Oaxaca persisting 23 years, spreading, held in a short, per-
and Chiapas; Guatemala: in the departments of El sistent basal sheath, 47(12)cm long, straight or
Quiche and Huehuetenango. slightly contorted, rigid, 12mm wide, often slightly
TDWG codes: 79 MXC-PU MXG-VC MXS-GR twisted, green or glaucous green; margins minutely
MXS-OA MXT-CI 80 GUA serrulate; apex acute-acuminate or mucronate; sto-
mata in fine grey-white lines on all surfaces. Pollen
Conservation cones short cylindrical, elongating to 22.5cm, yel-
low. Seed cones solitary or in whorls of 25, short
Although a valuable timber tree logged locally, the pedunculate, usually falling soon after seed disper-
main reason for its decline is deforestation and/or sal, ovoid-conical when closed, near-symmetri-
forest degradation. cal, (2)37cm long and 23cm wide, opening to
IUCN: EN [B2a (ii, iii, v)] 45cm wide. Seed scales thin woody, rigid, oblong,
spreading wide except basal scales, chestnut brown;
apophyses nearly flat to slightly raised, transversely
Pinus sylvestris L., Sp. Pl. 2: 1001. 1753. keeled, more or less rugose, ripening from green
to pale grey-brown or reddish brown; umbo small,
Etymology obtuse, unarmed or sometimes with a small, brittle
prickle. Seeds ovoid-oblong, 35mm long, grey-
The Latin species epithet sylvestris means growing brown with dark specks, or dark brown entirely;
in the forest. wing 1015mm long, 56mm wide, light brown or
grey-brown.
Vernacular names
Taxonomic notes
Scots pine; pin sylvestre (French); Gemeine Kiefer
(German); furu (Norwegian); tall (Swedish); sosna As can be expected, this is a variable species and
zwyczajna (Polish); sosna lesnaya, sosna obiknoven- more than 50 botanical varieties or subspecies have
naya (Russian) been named and described. Only three (including
var. sylvestris) are recognized here.
Description
Distribution
Trees to 35(40)m tall (most trees in natural habi-
tats not taller than 25m); trunk to 1.5m d.b.h., Eurasia: from N Spain and Scotland in the west to
straight or slightly crooked, sometimes forked in the Russian Far East, from Lapland in the north to
crown. Bark thin and papery, flaking, orange-brown Turkey in the south.
on young trees and in the crown of older trees, TDWG codes: 10 FIN GRB NOR SWE 11 AUT-AU
becoming thick and scaly on lower trunk, break- AUT-LI BGM-BE BGM-LU CZE-CZ CZE-SL GER HUN
ing into irregular plates, purplish brown weather- POL SWI 12 FRA-FR POR SPA-AN SPA-SP 13 ALB BUL
GRC ITA-IT ITA-SM ROM TUE YUG-BH YUG-CR YUG- fibreboard, and various wood-based materials.
KO YUG-MA YUG-MN YUG-SE YUG-SL 14 BLR BLT-ES In Russia and Scandinavia resin is extracted by
BLT-KA BLT-LA BLT-LI KRY RUC RUE RUN RUS RUW destructive distillation from the stumps and roots
UKR-MO UKR-UK 30 ALT BRY CTA IRK KRA TVA WSB of felled trees to produce Stockholm tar. In much
YAK 31 AMU KHA PRM 32 KAZ 33 TCS-AB TCS-GR of western Europe it is a widely planted forestry tree
34 TUR 36 CHI-NM CHM-HJ CHM-JL 37 MON for timber; it was introduced in the USA for simi-
lar purposes and for growing as Christmas trees.
Ecology Scots pine is or was also used to stabilize dunes, but
not those close to the sea as it is not very resistant
Across its enormous range Pinus sylvestris grows to salt-laden winds. In Belgium, the Netherlands,
naturally in a variety of habitats, the common Germany, and Denmark such plantations have led 789
denominator of which is deficiency of nutrients in to massive spontaneous spread of pines onto heath-
the soil. Thus on the Atlantic seaboard with high land and the last remaining inland dune areas,
levels of precipitation it occupies ancient igneous or and while the old plantations have in many places
metamorphic rocks with little or no soil in Scotland matured to mixed woodland now managed as mul-
and Norway up to 70 N, while south of the Baltic tiple use or even nature reserves, the invasivenes
Sea it grows on podzolized glacial sands left after onto Calluna heathland is seen as a menace to bio-
the Ice Age. In the central Alps it is restricted to the diversity and an ancient semi-natural landscape. In
drier slopes and valleys below other conifers like horticulture a large number of cultivars is known,
Larix and Picea, while in the Caucasus it ascends including dwarf forms from witches brooms; the
to 2600m on rocky outcrops and scree. In much of species is being planted as an amenity tree in many
Siberia it occupies the drier sites, but in Scandinavia countries.
and NE Europe it often borders acidic peat bogs. In
the steppes of Russia and Mongolia it occurs only 3 varieties are recognized:
along stream courses. Pinus sylvestris most com-
monly forms open pine forests and woodlands but
in many areas it is associated with conifers like Picea, Pinus sylvestris L. var. sylvestris. Type: [Habitat
Larix, Juniperus and with broad-leaved trees, espe- in Europae borealis sylvis glareosis], Illustration
cially Betula spp. and Populus tremula. In old growth Pinus sylvestris in Dalchamps, Hist. General. Pl.
stands there is often a well developed ground cover 1: 45. 1587. Fig. 249
of Vaccinium spp. or Empetrum nigrum in Atlantic
regions, and such pine forests are rich in mycorrhi- Pinus sylvestris L. var. scotica Beissn., in Jaeger &
zal fungi. Beissner, Ziergehlze, ed. 2: 488. 1884.
Uses Description
Scots pine is an important timber tree, but most of Young shoots usually rough with pulvini, light
the production goes to the paper industry. In the brown; bud scales (cataphylls) red-brown. Leaves
past it was more often put to use as mining props 57(8)cm long. Seed cones variable, but up to 7cm
and for interior construction; such uses are still long.
common in E Europe. Most of the pine used for fur-
niture in W Europe is actually spruce (Picea abies), Taxonomic notes
which has a smoother grain and is less resinous, but
often has more and darker knots, which are the There are no morphological or anatomical characters
discarded lower branches on the trunks of densely known to clearly distinguish the disjunct Scottish
planted trees. Other uses of Scots pine wood are (or population from those of northern Europe, and con-
were) street paving blocks, railway sleepers, fencing, sequently the Caledonian pine is here treated as
crates, pallets, boxes, laminated wood, particleboard, Pinus sylvestris var. sylvestris.
after shedding leaves, glabrous, new shoots yellowish often has arrested this succession in favour of
brown, pale brown or slightly glaucous, turning light the pines, which are of economic importance as a
brown to grey-brown or grey. Buds oblong, acute; forestry tree.
terminal bud to 20mm long; lateral buds smaller, all
slightly resinous; cataphylls appressed, pale brown. Uses
Leaves in fascicles of 2, sometimes 3, more or less
remote, persisting 23 years, spreading, held in a Chinese red pine is an important forestry tree yield-
1015mm long, persistent basal sheath, 615cm ing timber for construction; its wood is hard and
long, straight or curved, rigid, 11.5mm wide, often strong with a straight grain. The wood is used for
slightly twisted, dark green; margins minutely ser- mining props, railway sleepers, to build wooden
rulate; apex acute or acuminate; stomata in fine lines bridges and carts or wagons, to make tools, and for 791
on all surfaces. Pollen cones short cylindrical, ca. artificial fibres. The resin is extracted from the bark
2cm long, yellow. Seed cones solitary or in pairs, and leaves and produces turpentine; another prod-
short pedunculate, usually persisting a few years but uct of these parts is tannin used to make leather
opening soon, ovoid when closed, near-symmet- from hides. Essential oils distilled from the leaves
rical, 58.5cm long and 3.55cm wide, opening to and the pollen are used in traditional Chinese medi-
57.5cm wide. Seed scales woody, rigid, oblong to cine. In China, this species is commonly planted in
obovate, spreading wide except basal scales, often plantations for forestry purposes and occasionally
recurving, brown; apophyses prominently raised, as an ornamental tree. Elsewhere it is uncommon,
variously shaped, transversely keeled, ripening from even in Japan, despite its suitability as an amenity or
green to yellowish brown or pale brown; umbo pyra- forestry tree. While the existence of a few botanical
midal, armed with a sharp, curved prickle. Seeds varieties, some or all of which may also be in culti-
ovoid-oblong, 68 45mm, pale brown, mottled; vation, indicates its potential, no cultivars have been
wing 1218mm long, 57mm wide, light brown or selected and named.
grey-brown.
3 varieties are recognized:
Distribution
China: from Jilin and Liaoning in the NE to Yunnan Pinus tabuliformis Carrire var. tabuliformis. Type
in the SW and from Shandong in the E to Qinghai not designated.
and Sichuan in the W.
TDWG codes: 36 CHC-CQ CHC-SC CHC-YN Pinus tabuliformis Carrire var. brevifolia S. Y. Wang
CHI-NM CHI-NX CHM-LN CHN-BJ CHN-GS & C. L. Chang, Fl. Henan: 1: 135. 1981.
CHN-HB CHN-SA CHN-SD CHN-SX CHN-TJ CHQ
CHS-HN Description
Conservation Pinus taeda L., Sp. Pl. 2: 1000. 1753. Type: USA:
[Habitat in Virginiae, Canadae paludosis],
IUCN: LC J. Clayton 496 (lectotype BM).
Etymology
Pinus tabuliformis Carrire var. mukdensis (Uyeki
ex Nakai) Uyeki, J. Chsen Nat. Hist. Soc. 3: 45. The species epithet is a Latin word for pine tree or
1925. Pinus mukdensis Uyeki ex Nakai, Bot. Mag. its wood; its other meaning was a torch.
(Tokyo) 33: 195. 1919; Pinus tabuliformis Carrire
subsp. mukdensis (Uyeki ex Nakai) Businsk, Acta Vernacular names
792 Pruhoniciana 68: 26. 1999. Type: North Korea:
[locality not stated], H. Uyeki 2350 (holotype TI). Loblolly pine, Southern pine
Description Description
Bark dark grey, longitudinally or irregularly fissured. Trees to 45m tall (usually to ca. 30m); trunk to 1.6m
Leaves 11.2mm wide. Seed cones 48.5cm long d.b.h., usually a straight bole. Bark thin, slightly
and to 7cm wide when opened. scaly, orange-red in crown and upper bole, becom-
ing thicker, breaking into more or less square, scaly
Distribution plates on large trunks, darker red-brown. Branches
spreading wide, self-pruning leaving a clear bole,
NE China: Jilin (?), Liaoning; North Korea. forming a broadly conical or domed crown. Foliage
TDWG codes: 36 CHM-LN 38 KOR-NK branches moderately slender, 0.51cm thick, rough
with pulvini from fallen leaf fascicles, glabrous,
Conservation prominently ridged, orange-brown to reddish
brown, becoming darker with age. Buds cylindrical,
IUCN: LC acute, 1015(20)mm long, slightly resinous; cata-
phylls pale red-brown; margins fringed white; apex
reflexed. Leaves in remote fascicles of (2)3, held by
Pinus tabuliformis Carrire var. umbraculifera persistent, 1525mm long basal sheaths, remain-
T. N. Liou & Q. L. Wang, in Liou, Ill. Fl. Lign. Pl. ing 3 years on branchlet, spreading, straight, (10
N.E. China: 97, 548. 1958. Type: China: Hebei, )1220(23)cm long, rigid but more or less pliant,
Anshan Shi, Chin. coll. n.v. (holotype PE?). slightly twisted, 1.52mm wide, lustrous light green;
margins minutely serrulate; apex acute to abruptly
Description subulate; stomata in lines on all surfaces. Pollen
cones cylindrical, 24cm long, 710mm wide, radi-
Trunk monopodial only towards base, usually forked ally spreading, yellow. Seed cones solitary or in pairs
from below the crown. New shoots yellowish or pale (sometimes more) on short peduncles, opening and
brown, not or only slightly glaucous. Leaves rigid, falling soon, variable in size, (5)612(15)cm long,
1.21.5mm wide. narrowly conical to oblong when closed, mostly
symmetrical, opening to 69cm wide, becoming
Distribution more ovoid with a tapering, sometimes oblique
base. Seed scales thin woody, oblong, nearly flat,
China: Hebei (Central Liaoning, Anshan Shi). dull brown; apophyses slightly raised, those on basal
TDWG codes: 36 CHN-HB scales more gibbous, more or less rhombic or with
a rounded upper margin, transversely keeled, often
Conservation radially striated or wrinkled, dull pale brown; umbo
dorsal and central, pyramidal or recurved, terminat-
IUCN: LC ing in a strong, sharp prickle. Seeds obovoid, slightly
(5)1020(22)cm long, slender, pliant, slightly stunted. On the coastal slopes it comes down to
twisted, 0.71mm wide; margins serrulate; apex 600m a.s.l. Due to this altitudinal range, it occurs in
acute; stomata in fine lines on all surfaces. Pollen different forest zones from warm temperate to sub-
cones clustered, spirally arranged, short cylindrical, alpine, but at the lower and middle elevations it is
1.52cm long, yellow, becoming yellowish brown. mostly restricted to open spaces, exposed ridges and
Seed cones solitary or sometimes in pairs, persistent places with sandy, acidic and nutrient-poor soils. It is
on short peduncles, spreading or slightly reflexed, associated with various species of oak (Castanopsis,
(3)49(10)cm long, more or less asymmetrical, Quercus), forming pine-oak woodland in such
narrowly ovoid when closed, widening to 2.55cm places.
when open. Seed scales thin woody, rigid, oblong,
794 ca, 2.5 1.3cm at mid cone (in larger cones), straight Uses
when spreading, dull brown; apophyses rhombic in
outline or with rounded upper margin, nearly flat to Taiwan black pine has good quality timber with suit-
slightly raised, transversely keeled, slightly rugose, able strength for construction, e.g. in buildings and
lustrous brown; umbo broadly ellipsoid, flat, armed wooden bridges, and use as railway sleepers and
with a minute, often deciduous prickle or unarmed. mine props. Its wood is also used for building fences
Seeds ellipsoid-ovoid, 56mm long, slightly flat- and gates, crates and boxes, panelling, flooring, fur-
tened; wing 1520mm long, persistent. niture making, industrial and domestic woodware,
tools, plywood, fibreboard and wood pulp. In horti-
Taxonomic notes culture it is rare except for the use for bonsai grow-
ing, which is very popular in E Asia.
Pinus taiwanensis is similar to P. hwangshanensis,
P. luchuensis and P. densiflora and these species are 2 varieties are recognized:
closely related according to phylogenetic analy-
ses using DNA sequence data. Recently, Businsk
(2003) revisited the morphology of some of these Pinus taiwanensis Hayata var. taiwanensis. Type:
pines and separated some trees in Taiwan as a new Taiwan: Central Mts., T. Kawakami & U. Mori 2097
species, Pinus fragilissima. Most of the characters (lectotype TI). Fig. 250
evaluated are either similar to those of P. taiwanen-
sis, or they show overlapping states. The seed scales Description
are described as thin in the formal description and
elsewhere as fragile but these are difficult to quan- Leaves (5)1015(17)cm long. Seed cones (3)4
tify and may be attributes of the other taxa as well. 6(8)cm long.
What remains of this are somewhat longer leaves and
only slightly longer seed cones. Species are expected Distribution
to show distinct character state differences, without
overlapping states. This taxon should be given the Taiwan.
status of a variety only, as will be done below. TDWG codes: 38 TAI
Distribution Conservation
Taiwan. IUCN: LC
TDWG codes: 38 TAI
Ecology
Pinus taiwanensis Hayata var. fragilissima out, reddish brown. Branches spreading or slightly
(Businsk) Farjon, comb. et stat. nov. Basionym: ascending, the higher order branches flexible but
Pinus fragilissima Businsk, Novon 13 (3): 282. not pendulous. Shoots rough, reddish brown, often
2003. Type: Taiwan: Taitung Co., 1 km N of Wulu glaucous. Cataphylls subulate, erose-ciliate at mar-
along Southern Cross-Island Hwy., R. Businsk gins, scarious, brown. Vegetative buds oval-oblong
32172 (holotype PR). to cylindrical; terminal bud 1520mm long; lateral
buds smaller, not resinous. Fascicle sheaths ini-
Description tially up to 25mm long, orange-brown, in mature
fascicles reduced to 1218mm, grey-brown. Leaves
Leaves (12)1620(22)cm long. Seed cones (5)6 in fascicles of 4(35), persisting 23 years, lax and
9(10)cm long; seed scales often somewhat thinner drooping but not pendant, (14)1618(25)cm long, 795
than in var. taiwanensis. 0.71(1.3)mm wide, serrulate at margins, acute,
bright green. Stomata on all faces of the leaves.
Distribution Pollen cones ovoid-oblong to cylindrical, 1.52cm
56mm when shedding pollen, yellowish. Seed
Taiwan (Taitung Co., Kuan Shan massif). cones subterminal, in whorls of 24, rarely solitary,
TDWG codes: 38 TAI on (15)2025mm long peduncles, semi-serotinous,
persisting 13 years after shedding seeds, falling with
Conservation peduncles. Mature cones ovoid to broadly ovoid,
nearly symmetrical or asymmetrical, (3.5)47(7.5)
IUCN: NT (3)3.56cm when open. Seed scales thin woody,
parting usually within 12 years after maturity,
oblong, straight or slightly curved. Apophysis raised,
Pinus tecunumanii Eguiluz & J. P. Perry, Revista Ci. transversely keeled, on proximal scales more or less
Forest. 8: 4. 1983. Pinus patula Schiede ex Schltdl. gibbous, striate, dull light brown or slightly lustrous.
& Cham. subsp. tecunumanii (Eguiluz & J. P. Perry) Umbo dorsal, flat or slightly raised, with a minute,
Styles, F.A.O. Forest Genet. Resources Inform. 13: deciduous prickle, grey. Seeds obliquely ovoid, 47
50. 1984. Type: Guatemala: Baja Verapaz, Sierra de 24mm, dark grey-brown, with blackish dots, or
Chuacus, San Jronimo, T. Eguiluz 2 (holotype GH). blackish grey; wings articulate, 1013 48mm,
grey-brown.
Pinus oocarpa Schiede ex Schltdl. var. ochoterenae
Martnez, Anales Inst. Biol. Univ. Nac. Mxico 11: 65. Distribution
1940.
S. Mexico: Oaxaca, Chiapas; Belize; Guatemala;
Etymology Honduras; El Salvador; Nicaragua.
TDWG codes: 79 MXS-OA MXT-CI 80 BLZ ELS GUA
This species was named after Tecun Uman, a leader HON NIC
of the Quiche Indians who was killed during the
Spanish conquest of the Central American Isthmus. Ecology
been however greatly accelerated by humans. Here, Pinus teocote Schiede ex Schltdl. & Cham., Linnaea
open pine stands with grasses, Pteridium aquilinum, 5: 76. 1830. Type: Mexico: Veracruz, Pico de
Rubus, Calliandra, and Leucaena are predominant as Orizaba, (Volcan Citlaltepetl), C. J. W. Schiede &
long as the disturbances do not lead to further degra- F. Deppe s.n. (holotype HAL).
dation. In less disturbed areas, mostly at higher alti-
tudes, P. tecunumanii is often associated with other Etymology
pines, such as P. oocarpa, P. maximinoi, and P. pseu-
dostrobus, and at the more mesic sites P. ayacahuite Teocote was apparently one of the Aztec names used
and P. strobus var. chiapensis. Abies guatemalensis and for pines in Mexico.
Cupressus lusitanica are other conifers on these high
796 mountain ridges. On the Atlantic slopes in Chiapas Vernacular names
a mixed angiosperm forest with Liquidambar,
Magnolia, Clethra, Carpinus, Symplocos, Quercus and Aztec pine, Teocote pine; pino chino, pino colorado,
many other species predominates, and Pinus tecu- pino real, pino rosillo (Spanish)
numanii and other pines occur either on poorer sites
or at an earlier stage in a sere leading back to domi- Description
nance of broad-leaved trees.
Trees to 2025m tall, d.b.h. to 75cm; trunk mono-
Conservation podial, sometimes forked, straight. Bark thick,
rough and scaly, forming longitudinal plates
While occasionally still abundant and of very tall divided by deep, wide fissures, dark greyish brown.
stature, this pine is now usually scattered in small, Branches spreading horizontally or curved down;
disjunct populations and has been depleted by over- branches of higher orders slightly pendulous, form-
exploitation and forest clearing, especially at lower ing a pyramidal to rounded crown. Shoots smooth,
elevations, to such an extend that many of these pop- orange-brown. Cataphylls subulate, curved at
ulations are now vulnerable to extinction (Dvorak & apex, with ciliate margins, brown. Vegetative buds
Donahue, 1992). ovoid-oblong; terminal bud 1015mm long; lateral
IUCN: VU [A2cd, A3cd; B2ab (ii, v)] buds ovoid-acute, smaller, all not resinous. Fascicle
sheaths initially up to 20mm long, in mature fascicles
Uses reduced to ca. 10mm. Leaves in fascicles of 3(25),
persisting 23 years, straight or slightly curved, rigid,
Pinus tecunumanii is an important timber tree in (7)1015(18)cm long, 11.4mm wide, with serru-
Central America, where it can grow a straight bole late margins, acute-pungent, light green. Stomata
with large dimensions. It is largely exploited for sawn on all faces of leaves. Pollen cones ovoid-oblong to
timber and other local wood products in its native cylindrical, 11.8cm long, ca. 5mm wide, yellow-
range; potentials for wood pulp production are con- ish green. Seed cones commonly opposite, some-
sidered to be high if it was to be grown extensively times 13, on short, stout, curved peduncles falling
in plantations. This taxon has received considerable with cones. Mature cones ovoid to ovoid-oblong
interest from foresters as a species for potential planta- when closed, slightly asymmetrical, with a broad,
tion forestry to be introduced in tropical countries. A flattened but oblique base when opened, (3)46(
comprehensive collection of seed and specimens was 7) 2.55cm when open, often persistent. Seed
carried out by the Oxford Forestry Institute (OFI), as scales thick woody, oblong, straight or recurved.
well as by other organizations, throughout its entire Apophysis slightly raised, in some cones more or
range. A major limitation to introductions on a large less flat, transversely keeled, apical margin angular
scale is the limited availability of seeds, both from or crenate, light brown, in some cones with radial
natural stands and from so-called seed orchards. It marks. Umbo dorsal, flat to blunt-pyramidal, with a
has been planted as a forestry tree in Africa, India, minute, deciduous prickle. Seeds 35mm long, dark
South America and Australia (Queensland). It is not grey-brown; 1218 68mm, translucent, yellowish
known to be used as an amenity tree. with a dark tinge.
peduncles, falling shortly after seed dispersal, ovoid- lize sand dunes in coastal areas near urbanization. It
conical when closed, 46(7)cm long, opening to is also widely planted as an ornamental tree in Japan
(broadly) ovoid, 34.5cm wide. Seed scales woody, and Korea (somewhat less commonly in other coun-
rigid, oblong; apophyses nearly flat to slightly raised, tries) and some cultivars have been selected especially
transversely keeled, rhombic or with a rounded upper in Japan to suit traditional Japanese gardening. It is
margin, light brown, more or less lustrous; umbo dor- also being used in bonsai growing. In the USA this
sal, small, unarmed. Seeds obovoid, slightly flattened, species was widely planted for afforestation in coastal
57mm long, grey-brown; wing oblong, 1015mm areas of New England, until susceptibility to pests and
long, pale brown with dark stripes. diseases put an end to these schemes and forced the
species back to arboreta and parks, where solitary
798 Distribution trees are usually safe. Minor uses in Korea are of the
needles in pastry and in (medicinal) soft drinks.
Japan: S Hokkaido, Honshu, Kyushu, Shikoku; South
Korea (near the coast).
TDWG codes: 38 JAP-HK JAP-HN JAP-KY JAP-SH Pinus torreyana Parry ex Carrire, Trait Gn.
KOR-SK Conif.: 326. 1855.
Ecology Etymology
This is a species of pine growing at low to middle This species was named by W. E. Parry, who failed to
elevations (up to ca. 1000m above sea level) in the validly publish it, after the American botanist John
coastal hills and mountains of the islands of Japan Torrey (17961873).
and South Korea, where the climate is warm temper-
ate (with little or no frost) and moist. These regions Vernacular names
would have had a predominantly deciduous angio-
sperm forest cover, with conifers mixed in especially Torrey pine, Soledad pine, Del Mar pine
on poor, water-logged soils and on dry slopes and
mountain ridges. Pinus thunbergii would have occu- Description
pied these habitats as well as those in close proximity
to the sea coast. Extensive cultivation has removed Trees to 15(23)m tall; trunk to 1m d.b.h., mostly
the natural vegetation in most areas, but as a pioneer curved or crooked in habitat. Bark rough and scaly,
species P. thunbergii has been able to hold its own; deeply fissured on trunk, reddish brown turning to
it has been much planted in afforestation schemes purplish grey. Branches spreading and assurgent or
from where it could spread in adjacent uncultivated ascending, irregular, forming an open or flattened
areas. Its tolerance of salty winds makes it a species crown. Foliage branches stout, ca. 10 mm thick (20 mm
that grows well on the sea coasts of Japan, both natu- thick on coning shoots), new shoots glabrous,
rally and when planted; naturally its trunk becomes greenish, becoming purplish brown to nearly black.
bent and the crown flattened under severe exposure. Buds ovoid-conical, 2030mm long, without resin;
cataphylls light brown with white fringed margins.
Conservation Leaves in fascicles of 5, rarely 4 or 6, held in up to
20mm long but later shortening, persistent basal
IUCN: LC sheaths, persisting 34 years, spreading, rigid, 1525 cm
long, straight or curved, slightly twisted, ca. 2mm
Uses wide, greyish green; margins minutely serrulate; apex
abruptly acute; stomata in lines on all faces. Pollen
The wood of this pine is similar to that of the Black cones ovoid-cylindrical, 23cm long, 810mm wide,
pine (Pinus nigra) and is used for general construc- yellow. Seed cones solitary or rarely in pairs on stout,
tion, poles, railway sleepers, fences, pallets and crates, 34cm long peduncles, maturing in 3 years, persist-
flooring, fibreboard, and wood pulp. Japanese black ing to 5 years, massive, 1016cm long, broadly ovoid
pine is mostly used as a windbreak tree and to stabi- to ovoid-conical, opening slowly and only partially
799
Plate 34. Pinus torreyana subsp. torreyana. 1. Habit of tree. 2. Leaves. 3. Seed cone. 4. Seed.
to 817cm wide, strongly resinous. Seed scales Pinus torreyana Parry ex Carrire subsp. torreyana.
thick woody, rigid, cuneate, with 2 deep seed cavi- Type: USA: California, San Diego Co., Torrey Pines
ties adaxially; apophyses strongly developed, mark- State Park, J. R. Haller 10450 (neotype UCSB).
edly raised, multi-angular or more or less rhombic, Pl. 34
sharply transversely keeled or with 45 convergent
ridges, lustrous light or red-brown; umbo dorsal Description
and central, pyramidal or slightly curved, with an
obtuse, hard apex. Seeds large, obovoid, slightly flat- Trees with erect trunks commonly 15, sometimes
tened, 1724mm long, 1014mm wide, light brown more than 20m tall. Seed cones ovoid when closed,
or darker mottled; wing reduced, up to 10mm long, less than 12cm wide. Seeds more or less evenly
800 surrounding the seed like a partial ring, deciduous brown, not or only lightly mottled.
and sometimes remaining attached to the seed scale.
Distribution
Distribution
USA: S California (San Diego Co., on the coast N of
USA: S California (San Diego and Santa Barbara San Diego).
Co.). TDWG codes: 76 CAL
TDWG codes: 76 CAL
Conservation
Ecology
The small population on the mainland that consti-
Pinus torreyana is a relict species now confined to lit- tutes this subspecies is in part (southern subpopu-
toral habitat on the coast (up to 1.6 km inland) and on lation) legally protected in the Torrey Pines State
two small islands off the coast of southern California. Park. However, the small overall size, fewer than
It grows from immediately above the high tide mark 3500mature trees covering ca. 320 ha in two sub-
to about 180m a.s.l. on rocky or sandy slopes. On populations, and close proximity to major urban
these sites it seems dependent on the daily fog that development, put the subspecies highly at risk of
comes in from the ocean in the afternoon, mitigating destructive events such as fires, pest epidemics and
the heat of the sun and the resulting excessive evapo- diseases. Trees outside the reserve are often not pro-
transpiration. It grows with a sparse chaparral and tected from development; they are sometimes incor-
few other trees; in ravines sometimes accompanied porated in urban landscaping and sometimes felled
by a few oaks (Quercus sp.) and Arbutus menziesii. (personal obs. 1992).
IUCN: CR [B2ab (ii, iii, v)]
Uses
Torrey pine is not used as a timber tree; at present Pinus torreyana Parry ex Carrire subsp. insu-
the two disjunct populations are protected by law. It laris J. R. Haller, Syst. Bot. 11 (1): 45. 1986. Pinus
is in cultivation in California in gardens and some torreyana Parry ex Carrire var. insularis (J. R.
arboreta, but rare elsewhere. In the better growing Haller) Silba, Phytologia 68: 64. 1990. Type: USA:
conditions of gardens it can grow to a large tree; a California, Santa Barbara Co., Santa Rosa Island,
specimen in New Zealand was 45m tall with a d.b.h. J. R. Haller 10448 (neotype UCSB).
of 1.5m in 1982 (Grimshaw & Bayton, 2009: 626).
Although its conservation seems more or less cared Description
fore at present, growing this species more widely as
an ex situ backup is to be recommended; it is also Stunted trees commonly less than 10m tall, with a
an interesting species to grow and requires a mild flat crown (due to severe growing conditions). Seed
climate with warm, sunny summers and (near) cones broad when closed, usually over 12cm wide,
absence of frost in winter. with strong, nearly straight umbos. Seeds with dark
spots.
2 subspecies are recognized:
This is a pine of wet moors in W European mountains Virginia pine, Scrub pine
and typical for the association Pinetum uncinatae
Kastn., Flssn. et Uhlig (= Vaccinio uliginosi-Muge- Description
tum Oberdorfer). This association has only Pinus
uncinata as a characteristic species, but it is usually Trees to 1518m tall, rarely to 25m; trunk to 50cm
accompanied by various species of Vaccinium, by d.b.h., straight or contorted, forked, or shrubby.
Calluna vulgaris, Eriophorum vaginatum, and some Bark thin and scaly, irregularly fissured and ridged
oligotraphent species of Sphagnum like S. recurvum on lower part of larger trunks, grey-brown, becom-
and S. magellanicum, most of these with a high pres- ing reddish higher up in the tree. Branches numer- 803
ence in the vegetation. Pinus uncinata occurs natu- ous, irregularly spreading and ascending, forming
rally at altitudes of between 600 and 1600m a.s.l. In an irregular, often dense, rounded or flat-topped
the Alps and Pyrenees it grows on moist slopes up crown. Foliage branches slender, rough with pulvini
to the tree line; in the Alps it is confined to the west- from fallen leaf fascicles, new shoots glabrous, pur-
ern and northwestern ranges that are much wetter plish brown, often glaucous, becoming red-brown
than the central and southern Alps. Here it some- to grey. Buds short and broad, ovoid, 510mm long,
times grows with Picea abies or Pinus sylvestris; in shortly acute, resinous or not; cataphylls red-brown,
the Pyrenees the spruce is naturally absent. with whitish fringed margins. Leaves in fascicles of 2,
held by short, persistent sheaths, remaining 34 years
Conservation on branches, spreading, rigid, 38cm long, often
strongly twisted, 11.5mm wide, light or dark green
IUCN: LC or yellowish green; margins minutely serrulate; apex
narrowly acute; stomata in inconspicuous lines on all
Uses surfaces. Pollen cones ellipsoid-cylindric, 1015mm
long, red-brown turning yellow at anthesis. Seed
Mountain pine is not commercially important; its cones solitary or in pairs, nearly sessile or on 1cm
main value is ecological and it has some significance long, tenacious peduncles, persistent but opening
as an ornamental tree. The wood is used as fuel and soon, spreading out or curved back, 3.57cm long,
is suitable for light construction, but as this species slightly oblique to symmetrical, conical when closed,
grows slowly and remains small the yield is limited. ovoid when open. Seed scales woody, rigid, oblong,
Its only commercial use is located in the Pyrenees, dull red-brown with a purple-red sealing band;
where fairly extensive stands exist. Some of the wood apophyses prominently raised or slightly thickened,
is excellent for special uses like turnery, woodware transversely keeled, light brown; umbo dorsal and
and musical instruments due to its relative density central, pyramidal, armed with a slender, strong, to
and hardness. It is uncommon in cultivation and 5mm long, straight or curved prickle. Seeds obovoid,
restricted to local parks and further afield to some slightly flattened, 46(7)mm long, pale brown with
botanic gardens and arboreta. darker spots; wing 1620mm long, narrow.
Distribution
Pinus virginiana Mill., Gard. Dict., ed. 8: Pinus
No. 9. 1768. Type not designated. Fig. 251 E USA: from New York State in the north to N
Alabama and Mississippi in the south-west and
Etymology South Carolina in the south of its range.
TDWG codes: 75 INI NWJ NWY OHI PEN WVA 78
This species name indicates its native origin, ALA DEL GEO KTY MRY MSI NCA SCA TEN VRG
Virginia, with which much of the eastern USA was
designated in colonial times.
to the terminal, small, incurved, obtuse-triangular in plantations and several hybrids with related spe-
umbo, lustrous light brown with darker umbo. Seeds cies have been established with timber production
obliquely obovoid, (3)59mm long, 3.55mm wide, in mind (e.g. the cross between P. strobus and P. wal-
yellowish brown or brown; wing (10)2035mm lichiana = P. schwerinii Fitschen). Bhutan pine is a
long, (5)810mm wide, grey-brown. widely used amenity tree and a number of cultivars
have been selected and are in the trade.
Distribution
2 varieties are recognized:
Himalaya: from Afghanistan (Hindu Kush) to
NE India, SE Xizang [Tibet] and NW Yunnan;
N Myanmar [Burma]. Pinus wallichiana A. B. Jacks. var. wallichiana. 805
TDWG codes: 34 AFG 36 CHC-YN CHT 40 EHM-AP Type: Illustration in Lambert, Descr. Pinus 2, t. 3
EHM-BH EHM-DJ EHM-SI NEP PAK WHM-HP WHM- Pinus excelsa. 1824 (lectotype designated here).
JK WHM-UT 41 MYA Fig. 252
Taxonomic notes with stomatal lines on the two adaxial surfaces; mar-
gins minutely serrulate. Pollen cones in small clus-
This apparently high altitude form from NE ters, short cylindrical. Seed cones variable in size and
India in an area also claimed by China as lying in shape, from short ovoid to long cylindrical, initially
Xizang [Tibet], see Flora of China 4: 24 (1999), was erect on stout peduncles, becoming curved down to
described from a type specimen only. A specimen pendulous, 410(15)cm long, solitary or with 23
in PE (Ying & Hong 759) from Tangmai in Xizang together. Seed scales soft woody, more or less flex-
[Tibet] appears to belong to it as well. ible at base, cuneate to oblong; apophyses rhombic
to oblong (at base and apex of cone), curved or more
Distribution or less straight, not recurved or more commonly
806 recurved near cone base, ripening to yellowish brown
NE India: Arunachal Pradesh (Assam, Kameng or dark brown, weathering grey-brown; apex thin and
District); China: SE Xizang [Tibet]. straight or slightly incurved, rarely upcurved; umbo
TDWG codes: 36 CHT 40 EHM-AP terminal, small and sunken or obtuse. Seeds obovoid
or ellipsoid, 810mm long, ca. 6mm wide, usu-
Conservation ally with a well developed wing ca. 16mm long and
7mm wide.
IUCN: LC
Taxonomic notes
Pinus wangii Hu & W. C. Cheng, Bull. Fan Mem. Pinus wangii has been accepted as a distinct species
Inst. Biol., ser. 2, 1 (2): 191. 1948. Type: China: in Flora of China 4 (1999) and in the World Checklist
Yunnan, Xichou Xian, Fadou, C. W. Wang 85830 and Bibliography of Conifers (Farjon, 1998, [2001])
(holotype KUN). but was only tentatively accepted by Farjon (2005b) in
the description of P. fenzeliana Hand.-Mazz. A com-
Etymology plex of closely related taxa (among which also feature
P. kwangtungensis and P. dalatensis) occurs in SW
The species epithet commemorates the Chinese bot- China and Viet Nam. The Czech botanist R. Businsk
anist C. W. Wang, who collected the type specimen (2008) has treated these taxa in a narrowly circum-
in SE Yunnan, China. scribed concept of species and subspecies, recogniz-
ing more species than most. Hu & Cheng (op. cit.)
Vernacular names compared their new species to P. parviflora of Japan,
but we now consider that species more distantly
Wang pine; mao zhi wu zhen song (Chinese) related. The morphology is highly variable in some
of these species and may not be sufficiently clear to
Description resolve the taxonomy; additional DNA analysis may
be needed, but could only help to resolve this spe-
Trees to 20m tall; trunk to 60cm d.b.h. Bark on young cies problem if based on sufficiently wide sampling.
trees and branches smooth, thin, becoming scaly and Ideally, clades resolved from DNA data should corre-
flaking, brown, dark brown or grey-brown on trunks late with groupings based on distinct morphological
of larger trees. Branches spreading wide, forming character states, and only groups that are congruent
broad, umbrella-shaped or irregular, flat-topped on both criteria should be given species names.
crowns. Foliage branches slender; young shoots red-
brown, initially pubescent, glabrous or with remnants Distribution
of pubescence in grooves in the second to third year,
turning greyish brown. Buds ovoid to cylindrical, China: SE Yunnan (Malipo, Xichou, Maquan);
not resinous; cataphylls brown. Leaves in fascicles of Viet Nam (Mai Chou) [The taxonomic status in
5, held by deciduous sheaths of flimsy, brown scales, Viet Nam remains unclear, see Hiep et al., 2004)]. It
spreading or curved slightly towards shoot, 2.56cm straddles the border between the two countries.
long, pliant, straight or curved, 11.5mm wide, green, TDWG codes: 36 CHC-YN 41 VIE
high mountain slopes where it forms extensive for- Pinus insularis Endl. var. tenuifolia (W. C. Cheng &
ests. It is most abundant on dry and sunny slopes Y. W. Law) Silba, Phytologia 68: 51. 1990.
and it can locally reach to the tree line, forming an
alpine shrub. It is adapted to survive on sites with Description
infertile and shallow, rocky soils exposed to erosion.
At high altitudes it can withstand severe frosts. It is Trees to 30m tall. Leaves not or slightly pendulous,
most commonly gregarious in pure stands, but can 1020(30)cm long, with 46 resin ducts. Seed cones
be found growing with other pines, e.g. P. armandii, opening at maturity, falling after seed dispersal.
P. tabuliformis and P. kesiya, or with Keteleeria eve-
lyniana, as well as with angiosperm trees and shrubs. Distribution
808
Uses SW China: S Sichuan, Yunnan, E Guizhou, E Guangxi.
TDWG codes: 36 CHC-GZ CHC-SC CHC-YN CHS-GX
Yunnan pine is of considerable economic importance
in China, with natural stands covering between 5.5 Conservation
and 6million ha and 275,000 ha in plantations. Its
wood is used for fuel, as round wood for transmission IUCN: LC
poles and stakes, as sawn timber for construction,
both exterior and interior, for fences and gates, pal-
lets, crates and boxes, cooperage and vats, furniture, Pinus yunnanensis Franch. var. pygmaea (J. R. Xue)
veneers, all kinds of plywood and boards, tool han- J. R. Xue, Fl. Reipubl. Pop. Sin. 7: 258. 1978.
dles, pulp for paper, and to make chemical products Pinus densata Mast. var. pygmaea J. R. Xue, Acta
like plastics and cellulose derivatives. Resin is tapped Phytotax. Sin. 13 (4): 85. 1975; Pinus tabuliformis
for naval stores, in particular turpentine, and the bark Carrire var. pygmaea (J. R. Xue) Silba, Phytologia
is used to extract tannins. The leaves (needle litter) 68: 63. 1990. Type: China: Yunnan [locality not
provide fodder for animals and are distilled for oils stated], Y. C. Hseh 156 (holotype PE).
and medicinal products. This species is much planted
for wind breaks and erosion control. It is common in Description
China as an ornamental tree, but is not widely planted
in other countries, where it is mostly seen in arboreta A shrub-like high mountain form to 2m tall,
and other collections. It has a tendency to become branching from base. Leaves 713cm long, rigid, not
invasive as it is an aggressive pioneer species. drooping, with 23 resin ducts. Seed cones persis-
tent, more or less serotinous.
2 varieties are recognized:
Distribution
809
812
815
figure 288. Podocarpus figure 289. Podocarpus spinulosus on North Stradbroke Island, Queensland.
salignus foliage and young
seed cones
figure 296. Prumnopitys andina foliage and figure 297. Prumnopitys andina seed cones
pollen cones
816
Biota (D. Don) Endl., Syn. Conif.: 46. 1847, non Platycladus orientalis (L.) Franco, Portugaliae
Cassini (1825). Type: Biota orientalis (L.) Endl. Acta Biol., sr. B, Sist. Vol. Julio Henriques: 33.
[Platycladus orientalis (L.) Franco (Thuja orientalis 1949. Thuja orientalis L., Sp. Pl. 2: 1002. 1753. Type:
L.)] China: [locality unknown], leg. ign. LINN 1136.2
(lectotype LINN). Fig. 253, 254
Greek: platys = broad, flat; klados = branch; referring 817
to the flattened foliage branches. Thuja chengii Bordres & Gaussen, Bull. Soc. Hist.
Nat. Toulouse 73: 284. 1939; Platycladus chen-
Description gii (Bordres & Gaussen) A. V. Bobrov, 14. Symp.
Biodiv. Evol. Biol. Jena 1999: 18. 1999.
See the species description.
Etymology
Distribution
The species epithet means from the east, wherewith
As for the species. Linnaeus distinguished the species from his other
species in Thuja.
Taxonomic notes
Vernacular names
Detailed observations of the early stages of seed cone
development in both Microbiota and Platycladus Oriental Arbor-vitae, Chinese Arbor-vitae; ce bai
(Jagel & Sttzel, 2001) demonstrated that these taxa (Chinese)
are probably closely related. Both cones are quite
variable (variation is greater in Platycladus) and are Description
primarily distinct in size and in the number of bract-
scale complexes and ovules. Extremely reduced Trees to 2025m tall, evergreen, monoecious; trunk
cones of Platycladus resemble strongly developed monopodial or multistemmed from low above
cones of Microbiota; the latter seems merely a cone ground, to 22.5(4) m d.b.h., with stem(s) erect or
of Platycladus strongly reduced in size and number ascending. Bark on large stems becoming fibrous,
of parts, including seeds. Phylogenetic analyses of exfoliating in thin, long sheets, dull red-brown.
the genera in Cupressaceae (Brunsfeld et al., 1994; Branches spreading, ascending or erect, contorted in
Gadek et al., 2000) based primarily on molecular old trees, higher order branches mostly spreading,
data, confirmed the close relationship of these two forming a pyramidal, rounded or irregular crown.
taxa and their much greater distance from Thuja. A Foliage branches in flattened, more or less vertically
concise review of Platycladus, with botanical illus- disposed sprays, in older trees more horizontal or
trations, was published by Morgan (1999) in Curtiss irregularly disposed, persistent. Leaves decussate,
Botanical Magazine, hopefully with the effect that decurrent, scale-like, on lateral branchlets dimor-
this genus will be recognized by all as truly distinct phic, facials more or less rhombic to obtrullate,
from Thuja. with appressed, obtuse apex; laterals bilaterally flat-
tened, distal part spreading or reflexed, appressed or
free, with incurved, obtuse apex; leaves on ultimate
branchlets 1.52 11.5mm, with facials slightly
smaller than laterals; amphistomatic, stomata in
irregular lines; gland more or less linear, inactive chinensis and Populus tremula are followed in the
or sometimes active; leaf colour green or greyish succession by Quercus spp. (some of which may be
green. Pollen cones terminal on ultimate branchlets, evergreen) and Castanea. The most arid and steepest
solitary, subglobose, 23 2mm; microsporophylls slopes may only be covered with Pinus tabuliformis
(6)810(12), decussate, peltate, bearing 36 abax- and Platycladus, accompanied by Juniperus rigida
ial pollen sacs. Seed cones terminal, maturing in and other shrubs, and this vegetation may not rep-
one year to erect, ampulliform, glaucous or purplish resent a seral type, but an edaphically determined
cones 1520(25) 1018mm which open widely climax. The climate in NE China is above all char-
and turn brown when the seeds have ripened. Bract- acterized by very cold winters. As a pioneer of rela-
scale complexes 68, decussate, with mostly sterile tively dry, open vegetation on often unstable slopes,
818 upper pair, oblong to obtrullate, variable in size up P. orientalis has found abundant opportunity over
to 15 8mm, with a prominently recurved subapi- much of China, and even beyond (e.g. NE Iran), to
cal spine 38mm long, spreading wide, more or less establish itself and spread after introduction. It is
woody, thick, exposing often persistent seeds. Seeds much used in afforestation in NE and Central China
610(12?), ovoid to narrowly ovoid, 57 34mm, and commonly planted in Central Asia. Buddhism
greyish brown with a large, light brown hilum; wings has been instrumental in its spread especially in the
absent or rudimentary as two narrow strips near SW of China, where it can be abundant on steep
seed apex. slopes of river valleys, but is usually never very far
from settlements, monasteries or temples.
Distribution
Conservation
China: S Gansu, Hebei, Henan, Shaanxi, Shanxi, Nei
Mongol [Inner Mongolia]; Korea; Russian Far East. IUCN: NT
[This is possibly the most widely introduced cupres-
saceous conifer in Asia. In many areas inside and Uses
outside China it has escaped from cultivation and
established spontaneous populations. Only its natu- The wood of Oriental Arbor-vitae is used for build-
ral (indigenous) distribution is given here.] ing and construction of houses and temples where
TDWG codes: 31 AMU KHA 36 CHI-NM CHN-GS there are still trees of good size left. The foliage is in
CHN-HB CHN-SA CHN-SX CHS-HE 38 KOR-NK some parts of China much used for incense burning,
KOR-SK to which purpose the species has been introduced
widely outside its natural range. It is also one of the
Ecology most commonly planted amenity and ornamental
conifers, a tradition that goes back many centuries.
Platycladus orientalis is most probably a species of It is therefore a common tree in parks of towns and
the transitional open woodland zone between the cities in much of temperate Asia; it appears to be
steppes of Inner Mongolia and the deciduous oak, tolerant of drought as well as the air pollution that
oak-birch, and oak-pine forests of NE China. Even occurs within the urban environment. In some parts
within its natural range it is now almost invariably of China (e.g. Yunnan) and in other countries (e.g.
found in secondary vegetation or, nearest to its orig- Iran) it has probably been naturalized. In NE China
inal habitat, in more or less degraded woodland and it is being used in afforestation schemes on defor-
forest. As a pioneer species which is relatively long- ested hills and mountains. A substantial number of
lived, it can dominate certain slopes for a long time cultivars has been raised in Europe since its intro-
if further disturbances remain absent. Elsewhere duction to France around 1700, many of which are
it grows together with Pinus tabuliformis, less fre- now obsolete, while new cultivars continue to be
quently with P. armandii and Juniperus rigida; Betula selected and described.
Podocarpus LHr. ex Pers., Syn. Pl. 2 (2): 580. 1807 (nom. cons.).
Type: Podocarpus elongatus (Aiton) LHerit. ex Pers. [Taxus elongata Aiton]
(Podocarpaceae)
Distribution
Margbensonia A. V. Bobrov & Melikyan, Byull.
Moskovsk. Obshch. Isp. Prir., Otd. Biol. 103 (1): 59. Africa: S Nigeria, Cameroon, Angola; in E Africa
1998. Type: Margbensonia macrophylla (Thunb.) from extreme S Sudan to South Africa down to the
A. V. Bobrov & Melikjan [Podocarpus macrophyllus Cape; Madagascar. Asia: from E Nepal to Taiwan
(Thunb.) Sweet (Taxus macrophylla Thunb.)] and southern Japan; from Indochina throughout 819
Malesia (excluding Lesser Sunda Islands) to New
Greek: podo = foot, carpus = fruit; alluding to the Britain. Australia: SW corner of Western Australia,
receptacle and seed respectively of the female repro- Northern Territory (Arnhem Land), Queensland,
ductive organ (cone + seed). New South Wales, Victoria, Tasmania. SW Pacific:
Solomon Islands, New Caledonia, Vanuatu, Fiji,
Description Tonga, New Zealand. North America: S Mexico.
Caribbean Islands (excluding Bahamas). Central
(Decumbent) shrubs or more commonly trees, America (all countries). South America: Colombia,
dioecious or rarely monoecious, much branched. Venezuela, Guyana, Ecuador, Brazil, Bolivia,
Bark scaly or fibrous, peeling in vertical strips. NArgentina, S Chile.
Resin canals mostly in the leaves, none in the wood.
Primary branching in pseudo-whorls, plagiotropic Taxonomic notes
(Massarts model). Terminal buds distinctive, with
imbricate and/or spreading primary and secondary The taxonomy of the genus Podocarpus, after Pinus
scales. Leaves spirally inserted or subopposite on the most speciose (but not the most diverse) genus in
lateral foliage branchlets, relatively broad, bifacially the conifers, is perhaps the least well settled among
flattened, usually linear-lanceolate or linear-elliptic, the major coniferous genera. Whereas at the family
sessile to short petiolate, coriaceous, with a single level a limited number of phylogenetic studies have
raised, flat or sunken midrib and stomata in two now been published, which appear to mostly cor-
broad bands on abaxial side, rarely on both sides and roborate the narrower circumscription of the genus
then only few stomata on adaxial side. Pollen cones established in recent decades on morphological
axillary, solitary or clustered, sessile or on short grounds, no such studies have been attempted that
peduncles, each cone subtended by bud scales, flex- more or less sampled the genus Podocarpus compre-
ible and slender, catkin-like. Microsporophylls spi- hensively. There are two revisions of the genus based
rally attached to a slender rachis, triangular, with two on anatomy and/or morphology that need to be
basal pollen sacs containing bisaccate pollen. Seed briefly assessed here, in order to establish whether
cones axillary, pedunculate, solitary or rarely more they are useful for this Handbook. The first is by J. T.
together, consisting of 25 adnate bracts (upper 12 Buchholz and N. Gray and was published in a series
fertile); the sterile bracts fusing and greatly swelling of papers (variously authored by both or by Gray
to form a smooth, succulent, coloured receptacle. alone later) in the Journal of the Arnold Arboretum
Seeds 1, sometimes 2, obliquely attached to the apical in 19481962. The circumscription of Podocarpus
part of the receptacle, completely exposed, morpho- still included several other genera (as sections) and
logically inverted, drupe-like, completely covered only their section Eupodocarpus (an illegitimate
by a fleshy, green or sometimes reddish to bluish name under the Botanical Code) was equivalent to
epimatium. the genus Podocarpus as presently understood. This
section was divided into six subsections, informally
98 species named A-F. The emphasis was on a detailed study of
leaf anatomy, but the subsections were mostly defined 2a. Leaves 715mm long, 24mm wide, obovate
by geography and a few selected characters, indeed to oblanceolate (widest above the middle)
mostly anatomical. De Laubenfels (1985) followed P. humbertii
an earlier suggestion by Pilger (1926) that the pres- 2b. Leaves usually longer than 15mm (if smaller,
ence or absence of foliola (lanceolate bracts) below only 12mm wide), linear or widest below the
the receptacle could be used to subdivide the genus middle 3
Podocarpus (now in the narrower sense still in use) 3a. Leaves 736mm long, 12mm wide, narrowly
into two subgenera. Further special characteristics lanceolate to linear 4
then helped De Laubenfels to further subdivide the 3b. Leaves (1.5)218 cm long, 216 mm wide,
genus into 18 sections of from 110 species each elliptic-oblong to linear 5
820 and the two subgenera and 18 sections were then 4a. Seeds including the epimatium ca. 8 5mm.
formally named and described. In a phylogenetic Scales of terminal buds obtuse or acute. Leaves
context, it is unlikely that such a scheme, based as it 720(25)mm long P. perrieri
is on a few selected characters, would be supported 4b. Seeds including the epimatium ca. 14 8mm.
by a cladistic analysis of these and other anatomical Scales of terminal buds apiculate. Leaves
or morphological characters or of molecular data. (10)1530(36)mm long P. rostratus
Simply put, a different classification would result 5a. Leaves 25mm wide, up to 7.5cm long (mature
from the choice of different characters, and with- trees!). Bud scales elongated P. capuronii
out proper testing we do not know how artificial or 5b. Leaves 316mm wide, up to 18cm long (but
natural such a scheme might be. It must therefore highly variable!). Bud scales rounded to ovate
be concluded that neither of these two classifications P. madagascariensis
has been tested, either by repeated (phenetic or phy- 6a. Midrib on adaxial (upper) side of leaves dis-
letic) attempts using additional characters (as has tinct and continuously raised. Pollen cones
been the case with Abies and Pinus) or by a compre- always solitary P. milanjianus
hensive cladistic analysis of similar data or molec- 6b. Midrib on adaxial (upper) side of leaves indis-
ular data (as with Abies and especially Pinus). No tinct and/or fading towards the apex. Pollen
classification of Podocarpus is therefore presented in cones 25 together, rarely solitary 7
this Handbook. The large number of species and the 7a. Shrubs or small trees to 6m tall (occasionally
paucity of useful morphological characters make it to 20m). Leaves (3)45(7)mm wide
impractical to construct a single key to all species. P. elongatus
I therefore present keys to species of Podocarpus 7b. Trees to 3035 m tall (occasionally stunted
grouped on a geographical basis. An additional key on exposed mountain tops). Leaves (5)612
is presented to help identify species that are more or (18)mm wide 8
less commonly found in cultivation. 8a. Apex of leaves acute or acuminate. Pollen cones
(1)25 together. Receptacle below the seed
Key to the species of Podocarpus in Africa rudimentary; seeds including the epimatium
and Madagascar 1722mm long P. henkelii
8b. Apex of leaves obtuse or mucronate. Pollen
When using this key in the field only leaves on cones 12 together. Receptacle below the seed
branches of mature shrubs or trees, preferably from 814 812 mm, succulent; seeds including
sun-exposed foliage, should be taken into account. the epimatium 711mm long P. latifolius
Male (pollen) and female cones and seeds (they
occur on dioecious shrubs and trees) are described Key to the species of Podocarpus in China
as mature. (including Taiwan), NE India, Indochina,
and Japan
1a. Scales of terminal buds all imbricate or
incurved 2 When using this key in the field only leaves on
1b. Scales of terminal buds with at least some outer branches of mature trees, preferably from sun-
scales spreading or recurved 6 exposed foliage, should be taken into account. Male
(pollen) and female cones and seeds (they occur on 9a. Receptacles subtended by two 26 mm long
dioecious trees) are described as mature. bracts (foliola); epimatium around the seeds
without a crest. New leaves flushing red
1a. Buds globose to short conical, with imbricate, P. neriifolius
triangular scales 2 9b. Receptacles subtended by two 12 mm long
1b. Buds variously shaped but not globose, with bracts; epimatium around the seeds with a
free, erect to spreading or recurved (outer) crest. New leaves flushing yellowish green
cales 3 P. subtropicalis
2a. Adult leaves (3)510(12)cm long, lanceolate
to linear-lanceolate; midrib on adaxial (upper) Key to the species of Podocarpus in Malesia,
side of leaves acutely raised P. nakaii excluding Borneo and New Guinea, and 821
2b. Adult leaves 914(22)cm long, linear; midrib including the Andaman Islands (India)
on adaxial side of leaves obtusely raised
P. rumphii Malesia is the phytogeographic region as defined in
3a. Adult leaves 1.25(7) cm long. Bud scales Flora Malesiana, and includes Peninsular Malaysia
erect or slightly preading 4 and Singapore and all the islands of the archi-
3b. Adult leaves (2.5)412(15) cm long. Bud pelago as far as New Guinea and the main chain
scales spreading or recurved, sometimes more of the Solomon Islands. Species which occur only
or less erect 6 in Borneo or New Guinea including the Bismarck
4a. Adult leaves 2.54.5mm wide, not longer than Archipelago and/or the Solomon Islands are dealt
3.5cm P. chingianus with in two keys specific to these areas; species which
4b. Adult leaves (4)58(10) mm wide, often are common to two or more of these areas appear
longer than 3.5cm (maximum length 57cm) consequently in more than one key. When using this
5 key in the field only leaves on branches of mature
5a. Pollen cones solitary, sessile, 825 68mm. trees, preferably from sun-exposed foliage, should
Small trees 1.510m tall P. costalis be taken into account. Male (pollen) and female
5b. Pollen cones solitary, pedunculate, 1530 cones and seeds (they occur on dioecious trees) are
3mm. Shrubs or trees to 25m tall P. pilgeri described as mature.
6a. Bud scales caudate, recurved. Midrib on adax-
ial (upper) side of leaves obtusely raised 1a. Buds short conical or subglobose; bud scales
P. macrophyllus imbricate 2
6b. Bud scales triangular-lanceolate to lanceolate, 1b. Buds elongated, acute; outer bud scales free,
erect, spreading or sometimes recurved. Midrib erect or spreading 4
on adaxial side of leaves acutely raised 7 2a. Bud scales rounded at apex. Midrib of leaves on
7a. Buds 1.53mm long. Adult leaves from nearly adaxial (upper) side acutely raised, narrow, less
oval to linear, with an acute or obtuse apex than 1mm wide P. teysmannii
P. polystachyus 2b. Bud scales triangular, with an acute apex.
7b. Buds (2)35(8)mm long. Adult leaves lan- Midrib of leaves on adaxial side obtusely raised,
ceolate to linear-lanceolate, with an acute or 1mm wide 3
acuminate apex 8 3a. Buds up to 4 4 mm, short conical. Seeds
8a. Pollen cones 1525(30) 22.5 mm. including the epimatium ca. 10mm long
Receptacles when ripe 916 mm long; seeds P. laubenfelsii
including the epimatium ca. 9 7mm 3b. Buds (2.5)45(8) mm long, globose to
P. fasciculus short conical. Seeds including the epimatium
8b. Pollen cones 2040(60?) 23.5 mm. 1215mm long P. rumphii
Receptacles when ripe 712 mm long; seeds 4a. Adult leaves 0.83(3.5) cm long, 36 mm
including the epimatium (8)1015 810mm wide. Shrubs or small trees 5
9 4b. Adult leaves mostly longer. (Small) trees 6
5a. Adult leaves elliptic to obovate; apex obtuse to sometimes with a rounded apex. Receptacle
abruptly rounded P. glaucus red when ripe 14
5b. Adult leaves elliptic to oblong; apex acute to 14a. Pollen cones solitary, 6.58 mm wide. Small
obtuse P. lophatus trees as far as known P. palawanensis
6a. Adult leaves (4)58(10)mm wide 7 14b. Pollen cones solitary or more often with
6b. Adult leaves (6)818(20)mm wide 10 23(5) together, 24 mm wide. Potentially
7a. Buds 614 mm long; outer bud scales erect large trees 15
with recurved tips. Adult leaves 611cm long 15a. Midrib on adaxial (upper) side of leaves
P. atjehensis obtusely raised, fading towards the apex 16
7b. Buds 25 mm long; outer bud scales erect or 15b. Midrib on adaxial side of leaves acutely raised,
822 spreading, with straight tips. Adult leaves always continuous to the apex 17
(2)36(7)cm long 8 16a. Buds 24 mm long. Pollen cones to 25 mm
8a. Pollen cones solitary, sessile, 68 mm diam. long. Receptacles 1015mm long; seeds includ-
P. costalis ing the epimatium 1417 1013 mm
8b. Pollen cones solitary or more often with 23 P. macrocarpus
together, pedunculate when solitary, otherwise 16b. Buds 39 mm long. Pollen cones 5060
sessile, 2.53.5mm diam 9 (80?)mm long. Receptacles 8mm long; seeds
9a. Apices of adult leaves acute. Pollen cones soli- including the epimatium 811(13?) 78mm
tary or often with 23 together, sessile. P. levis
Receptacles 68 mm long when ripe; seeds 17a. Buds 1.53mm long. Pollen cones in clusters
including the epimatium 89 56 mm, red of 25, 1530mm long P. polystachyus
or purple P. rubens 17b. Buds 212 mm long. Pollen cones solitary
9b. Apices of adult leaves of various shapes, but or with 23 together, 2550(60)mm long
often obtuse to rounded. Pollen cones solitary, 18
pedunculate. Receptacles 1022mm long when 18a. Adult leaves 37(9)cm long, with an obtuse
ripe; seeds including the epimatium 1012 or rounded apex. Receptacles 6mm long; seeds
89mm P. pilgeri including the epimatium 6 5 mm, purple
10a. Adult leaves (1.5)2.55 cm long, elliptic to P. borneensis
obovate, with an obtuse or rounded apex 18b. Adult leaves (4)812(15) cm long, with an
P. ramosii acute apex. Receptacles 814 mm long; seeds
10b. Adult leaves (2.5)415(18.5)cm long, nearly including the epimatium (8)1015 78mm
oval to linear, rarely elliptic, with an acute, 19
obtuse or rounded apex 11 19a. Buds 25mm long, sometimes with a few lon-
11a. Leaves linear, deflected at the petiolate base ger outer scales to 8 mm. Receptacles when
P. deflexus ripe 810mm long P. neriifolius
11b. Leaves nearly oval to linear-lanceolate or linear, 19b. Buds 512 mm long, with all or most of the
not deflected 12 outer scales the same length. Receptacles when
12a. Adult leaves (8)1220 mm wide (510 cm ripe 1014mm long P. bracteatus
long), with obtuse or rounded apex
P. spathoides Key to the species of Podocarpus in Borneo
12b. Adult leaves 615(18)mm wide, with acute or
obtuse apex (if wider than 14mm, apex acute) When using this key in the field only leaves on
13 branches of mature trees, preferably from sun-
13a. Midrib on adaxial (upper) side of leaves indis- exposed foliage, should be taken into account. Male
tinct, narrow; adult leaves very gradually taper- (pollen) and female cones and seeds (they occur on
ing to a narrowly acute apex. Receptacle pink dioecious trees) are described as mature.
when ripe P. ridleyi
13b. Midrib on adaxial side of leaves distinct, acutely 1a. Bud scales imbricate 2
or obtusely raised; adult leaves acute or obtuse, 1b. Bud scales free, erect or spreading 5
2a. Buds 25mm long, scales rounded 3 11b. Bud scales narrowly triangular to lanceolate-
2b. Buds 48mm long, scales triangular 4 linear or long acuminate, erect or spreading.
3a. Buds 23 mm long. Leaves (2.5)48(9) cm Leaves linear-lanceolate, shortest ones lanceo-
long, (7)915mm wide P. globulus late or nearly oval 12
3b. Buds 35 mm long. Leaves 714 cm long, 12a. Bud scales long acuminate, erect. Pollen cones
(10)1217(20)mm wide P. teysmannii solitary, rarely in pairs. Receptacle with a single
4a. Bud scales with recurved apex. Pollen cones basal bract 35mm long P. confertus
35 together, short pedunculate 12b. Bud scales triangular to lanceolate-linear, erect
P. laubenfelsii or spreading. Pollen cones solitary or more
4b. Bud scales entirely appressed. Pollen cones often with 25 together. Receptacle with 2 basal
(2)35 together, sessile P. rumphii bracts 1.56mm long 13 823
5a. Leaves 17 cm long, rarely to 9 cm long, 13a. Buds 1.53 mm long. Leaves nearly oval to
313mm wide; shorter leaves usually elliptical, linear-lanceolate. Pollen cones 25 together,
sometimes obovate; longer leaves oblong or 1530 3mm. Seeds including the epimatium
linear-lanceolate 6 79 57mm P. polystachyus
5b. Leaves (2.5)515(18)cm long, (5)620mm 13b. Buds (2)315 mm long. Leaves linear-
wide, usually linear-lanceolate, sometimes lin- lanceolate or lanceolate. Pollen cones solitary
ear-oblong; shorter leaves sometimes elliptical or 23 together, (25)3075(80?) 24mm.
or nearly oval 11 Seeds including the epimatium (6) 815
6a. Buds 25mm long 7 68mm 14
6b. Buds 410mm long 9 14a. Buds 615 mm long, scales long linear-
7a. Buds 23mm long. Leaves elliptical to obovate, lanceolate, erect or spreading. Leaf apex acumi-
713mm wide; apex obtuse or rounded nate or acute. Epimatium with a distal crest
P. ramosii P. micropedunculatus
7b. Buds 25 mm long. Leaves linear-elliptic to 14b. Buds (2)39 mm long, scales lanceolate or
linear-lanceolate, (4)58(10)mm wide; apex rarely triangular, spreading. Leaf apex acute
acute, obtuse, or rounded 8 or obtuse. Epimatium smooth without a distal
8a. Leaves flushing green. Pollen cones always crest 15
solitary, pedunculate. Receptacle when ripe 15a. Midrib on adaxial (upper) side of leaves
1022mm long P. pilgeri obtusely raised, often fading towards the acute
8b. Leaves flushing red. Pollen cones solitary or or obtuse apex. Seeds including the epimatium
often 23 together. Receptacle when ripe 811mm long P. levis
68mm long P. rubens 15b. Midrib on adaxial (upper) side of leaves usually
9a. Bud scales acuminate, spreading. Pollen cones acutely raised, continuous to the acute apex.
solitary, rarely in pairs, ca. 15 34mm Seeds including the epimatium (8)1015mm
P. gibbsiae long P. neriifolius
9b. Bud scales triangular or lanceolate-linear, erect
or spreading. Pollen cones solitary or often 23 Key to the species of Podocarpus
together, 2050 2.55mm 10 in Papuasia
10a. Bud scales 46mm long, triangular, spreading.
Pollen cones solitary or in pairs, ca. 20 Papuasia is the phytogeographic region that includes
45mm P. brevifolius New Guinea, the Bismarck Archipelago and the
10b. Bud scales 410 mm long, lanceolate-linear, Solomon Islands (as on Plate 12 of the Times Atlas,
erect or slightly spreading. Pollen cones soli- 12th edition, but excluding the Santa Cruz Islands).
tary or with 23 together, 3550 2.53.5mm When using this key in the field only leaves on
P. borneensis branches of mature trees, preferably from sun-
11a. Bud scales triangular, spreading. Leaves linear- exposed foliage, should be taken into account. Male
oblong, shortest ones elliptical; apex obtuse or (pollen) and female cones and seeds (they occur on
rounded P. spathoides dioecious trees) are described as mature.
1a. Usually shrubs, only rarely (small) trees. Adult 9a. Pollen cones always solitary, 3040 68mm.
leaves not longer than 3cm 2 Buds 35mm long, with triangular to lanceo-
1b. Trees (can be stunted at high altitudes). Adult late, slightly spreading scales. Seeds including
leaves usually longer than 3cm 3 the epimatium 56mm wide P. archboldii
2a. Buds 35 mm long. Pollen cones ca. 40 9b. Pollen cones often 23 together or solitary,
38 mm. Receptacles when ripe 1215 mm 50(80?) 23.5mm. Buds 39mm long, with
long, thick, purple or purple-black; seeds lanceolate, spreading outer scales. Seeds includ-
including the epimatium 1013 89 mm, ing the epimatium 78mm wide P. levis
without an apical crest P. brassii 10a. Buds 614 mm long; bud scales with a long,
2b. Buds 23 mm long. Pollen cones 1025 acuminate or caudate apex 11
824 67mm. Receptacles when ripe 68mm long, 10b. Buds 1.57(8)mm long; bud scales triangular
purple; seeds including the epimatium 67 to lanceolate 13
5mm, with a small apical crest P. glaucus 11a. Adult leaves (11)1525mm wide. Receptacles
3a. Adult leaves (1.5)2.56cm long 4 when ripe 1016mm long P. ledermannii
3b. Adult leaves mostly longer than 5 cm, only a 11b. Adult leaves (5)712 mm wide. Receptacles
few sometimes shorter 5 when ripe 811mm long 12
4a. Pollen cones always solitary and pedunculate. 12a. All bud scales erect. Adult leaves (4)615cm
Receptacles when ripe 1022 mm long; seeds long, leaf apices acute P. pseudobracteatus
including the epimatium 1012 89 mm 12b. Outer bud scales spreading. Adult leaves
P. pilgeri 1025 cm long, leaf apices acute or obtuse
4b. Pollen cones solitary or often 23 together, P. salomoniensis
sessile. Receptacles when ripe 68 mm long; 13a. Buds on leading shoots (3)48mm wide and
seeds including the epimatium 89 56mm 47mm long, onion-shaped or truncate, with
P. rubens strongly recurved outer scales. Pollen cones
5a. Buds globose to conical, as wide as long, with 67mm wide P. crassigemma
imbricate scales. Leaves all linear P. rumphii 13b. Buds on leading shoots longer than wide, more
5b. Buds variously shaped but not globose, includ- or less conical, with erect or spreading outer
ing the free, erect or spreading outer scales lon- scales. Pollen cones 23.5mm wide... 14
ger than wide. Leaves variously shaped, not all 14a. Adult leaves 611 mm wide. Pollen cones
linear 6 always clustered with 25 together, 1530mm
6a. Leaves (strongly) deflected at the petiolate base long. Seeds including the epimatium 79mm
P. atjehensis long P. polystachyus
6b. Leaves spreading or sometimes drooping, not 14b. Adult leaves 815(18)mm wide. Pollen cones
deflected 7 solitary or often 23 together, 2540(50)mm
7a. Apex of adult leaves obtuse or rounded, leaves long. Seeds including the epimatium (8)10
510cm long, (8)1220mm wide. Receptacles 15mm long 15
when ripe 5mm long; seeds including the epi- 15a. Buds 25mm long, sometimes with a few lon-
matium 7 5mm, purple P. spathoides* ger outer scales to 8mm. New leaves flushing
7b. Apex of adult leaves acute, acuminate or obtuse, red P. neriifolius
never rounded, if adult leaves 20 mm wide 15b. Buds 23mm long, without longer outer scales.
(or wider), longer than 10 cm. Receptacles New leaves flushing pale whitish green
when ripe longer than 8mm; seeds including P. insularis
the epimatium larger than 7 5mm, variously
(purplish) green 8
8a. Midrib on adaxial (upper) side of leaves * Podocarpus spathoides has been found to be limited
obtusely raised, often fading towards apex 9 to Malaysia and trees from Papuasia that would
8b. Midrib on adaxial (upper) side of leaves acutely key out to belong here are not that species (Mill &
raised, continuous to apex 10 Whiting, 2012).
Key to the species of Podocarpus Key to the species of Podocarpus in
in Australia the SW Pacific
When using this key in the field only leaves on The SW Pacific for the purposes of this key includes
branches of mature trees, preferably from sun- New Caledonia, Vanuatu, Fiji, and Tonga. When
exposed foliage, should be taken into account. Male using this key in the field only leaves on branches
(pollen) and female cones and seeds (they occur on of mature trees, preferably from sun-exposed foli-
dioecious trees) are described as mature. age, should be taken into account. Male (pollen) and
female cones and seeds (they occur on dioecious
1a. Shrubs. Adult leaves 25mm wide 2 trees) are described as mature.
1b. Trees. Adult leaves (5)730mm wide 4 825
2a. Adult leaves 0.41.6cm long; leaf apices obtuse. 1a. Adult leaves 0.85cm long 2
Bud scales imbricate, broadly triangular 1b. Adult leaves (2.5)414(15)cm long 3
P. lawrencei 2a. Trees. Adult leaves 2.55 cm long, 69 mm
2b. Adult leaves 2.59(13) cm long; leaf apices wide P. affinis
acuminate or pungent. Bud scales free, (slightly) 2b. Shrubs. Adult leaves 0.82.2 cm long, 1.7
spreading, narrowly triangular 3 2.3mm wide P. gnidioides
3a. Apices of leaves apiculate, pungent; midrib on 3a. Buds small (usually ca. 2mm long); bud scales
adaxial (upper) side of leaves acutely raised. imbricate, obtuse or short triangular 4
Pollen cones 48mm long. Receptacles when 3b. Buds larger (usually more than 3 mm long);
ripe 610 67mm; seeds including the epi- bud scales free, erect, spreading or outer bud
matium 810 57mm P. spinulosus scales recurved, triangular-lanceolate, elon-
3b. Apices of leaves acuminate; midrib on adaxial gated or acuminate 6
side of leaves obtusely raised. Pollen cones 4a. Shrubs. Adult leaves 35(6) mm wide, with
(5)1020(22) mm long. Receptacles when a central groove on adaxial (upper) side
ripe 2025 1013 mm; seeds including P. novae-caledoniae
the epimatium 1220 812mm 4b. Trees. Adult leaves (5)718mm wide, with an
P. drouynianus obtusely raised midrib on the adaxial side 5
4a. Bud scales free, spreading. Adult leaves 5a. Adult leaves (9)1118mm wide. Seeds includ-
(13)1830mm wide, oval to oblong ing the epimatium without a distal crest
P. dispermus P. lucienii
4b. Bud scales imbricate, appressed. Adult leaves 5b. Adult leaves (5)711(13) mm wide. Seeds
(5)718(20) mm wide, linear to linear- including the epimatium with a (faint) crest
lanceolate 5 P. sylvestris
5a. Adult leaves (6)1025(30) cm long, linear 6a. Low, decumbent shrubs. Bud scales free, erect,
P. grayae not spreading. Adult leaves oblanceolate (wid-
5b. Adult leaves (3)511(15) cm long, linear- est beyond the middle) P. decumbens
lanceolate 6 6b. Erect shrubs or more often trees. Bud scales
6a. Buds 1.52.5 mm long. Midrib on adaxial free, spreading to recurved or reflexed. Adult
(upper) side of leaves obtusely raised but prom- leaves linear-lanceolate (widest at or below the
inent. Pollen cones 1020 23mm. Receptacles middle) 7
when ripe 1525(30) 1520mm, dark pur- 7a. Buds with distinctly elongated outer scales
ple or blue-black P. elatus 415mm long. Midrib on the adaxial (upper)
6b. Buds 3.58mm long. Midrib on adaxial side of side of leaves obtusely raised. Seeds including
leaves obscure. Pollen cones 3045 46mm. the epimatium 79mm long 8
Receptacles when ripe 57 5 mm, red 7b. Buds with triangular or lanceolate, sometimes
P. smithii acuminate outer scales 26(8) mm long.
Midrib on adaxial side of leaves acutely raised. 12.5(3) cm long, (1.5)23.5(4.5) mm
Seeds including epimatium (8)1015 mm wide, with a faint groove on the adaxial side
long 9 P. totara
8a. Pollen cones always solitary, 3.55 mm wide.
Receptacles when ripe 1013 89mm. Adult Key to the species of Podocarpus in Mexico,
leaves 68mm wide P. polyspermus Central America, and the Caribbean Islands
8b. Pollen cones with 23 together or sometimes
solitary, 2.53.5 mm wide. Receptacles when When using this key in the field only leaves on
ripe 8 45mm. Adult leaves 510(12)mm branches of mature trees, preferably from sun-
wide P. longefoliolatus exposed foliage, should be taken into account. Male
826 9a. Shrubs or small trees. Adult leaves 511 mm (pollen) and female cones and seeds (they occur on
wide. Pollen cones always solitary. Receptacles dioecious trees) are described as mature.
when ripe turning from red to blackish
P. pallidus 1a. Adult leaves (1)1.53(3.5)cm long, 35mm
9b. Trees. Adult leaves (6)815(18) mm wide. wide 2
Pollen cones solitary or more often with 23 1b. Adult leaves (1.5)215(20) cm long,
together. Receptacles when ripe red 10 (3)520(22)mm wide 3
10a. Buds 25mm long, sometimes with a few lon- 2a. Adult leaves narrowly lanceolate (widest below
ger outer scales to 8mm. New leaves flushing the middle), with apiculate-pungent or acute
red P. neriifolius apex. Small trees to 16 m tall (often only
10b. Buds 23mm long, without longer outer scales. 510m) P. urbanii
New leaves flushing pale whitish green 2b. Adult leaves oblanceolate to linear-oblanceo-
P. insularis late (widest above the middle), with acuminate
apex. Shrubs or small trees to 8m tall
Key to the species of Podocarpus P. ekmanii
in New Zealand 3a. Buds (on leading shoots) at least 5 mm long,
often twice as long, with ovate-apiculate to long
When using this key in the field only leaves on acuminate, more or less spreading (outer)
branches of mature trees, preferably from sun- scales 4
exposed foliage, should be taken into account. Male 3b. Buds (on leading shoots) not longer than
(pollen) and female cones and seeds (they occur on 5 mm, with ovate to triangular, rounded to
dioecious trees) are described as mature. apiculate or short acuminate, mostly imbricate
scales 6
1a. Shrubs. Pollen cones 12.5mm wide 2 4a. Adult leaves (7)1018(22)mm wide, lanceo-
1b. Trees. Pollen cones 38mm wide 3 late, with a grooved midvein on adaxial (upper)
2a. Adult leaves (0.5)0.82 cm long, 1.42 mm side P. costaricensis
wide, acicular, pungent. Receptacles when ripe 4b. Adult leaves 714 mm wide, linear-lanceolate
57mm long P. acutifolius to linear or narrowly lanceolate, with a raised
2b. Adult leaves 0.31 cm long, 22.5 mm wide, but fading midrib (midvein) on adaxial side
naviculate (boat-shaped) to ovate-linear, obtuse 5
or mucronate. Receptacles when ripe 810mm 5a. Pollen cones solitary or in pairs, 5060
long P. nivalis 34mm. Bud scales ovate-apiculate, free at apex
3a. Pollen cones solitary or more often in clusters P. coriaceus
of 25, 2030 58mm. Adult leaves (1.5)2 5b. Pollen cones always solitary, 3045 45mm.
3(3.5)cm long, 35(6.5)mm wide, with a nar- Bud scales long acuminate, more or less spread-
row but conspicuous groove on adaxial (upper) ing P. matudae
side P. cunninghamii 6a. Adult leaves 1.55.5 cm long, with an acute-
3b. Pollen cones solitary, sometimes in pairs, pungent or spinescent apex. Buds small, with
1215(20) 34 mm. Adult leaves (0.5) ovate or rounded scales 7
6b. Adult leaves (1.5)3.515(20) cm long, with 1a. Buds longer than wide at base, with outer scales
an acute, acuminate or obtuse apex (if pungent, 915mm long, acuminate or cuspidate, some-
adult leaves longer than 6 cm). Buds up to times elongated to become more or less leaf-
5mm long, with triangular or acuminate, rarely like 2
ovate or rounded scales 8 1b. Buds (usually) not longer than wide at base, with
7a. Receptacles when ripe 1015 mm long. outer scales up to 6mm long, of various shapes
Adult leaves 25.5 cm long, acute-pungent but with a short apex and not elongated 7
P. angustifolius 2a. Adult leaves (1.5)25 cm long; midvein on
7b. Receptacles when ripe 810 mm long. Adult adaxial (upper) side of leaves a continuous
leaves 1.54cm long, spinescent P. buchii groove 3
8a. Adult leaves (3)715(20)cm long, (12)15 2b. Adult leaves (2)413 cm long; midrib (mid- 827
20mm wide (juvenile leaves often substantially vein) slightly raised or obtusely raised, with
larger). Seeds including the epimatium or without a medial groove towards leaf apex
911mm long P. magnifolius 5
8b. Adult leaves (1.5)3.59 cm long, (5)6 3a. Adult leaves linear-lanceolate, 24(5) mm
14 mm wide. Seeds including the epimatium wide, strongly spinescent (with a hard, spine-
78mm long 9 like apex) P. glomeratus
9a. Midvein on adaxial (upper) side of leaves form- 3b. Adult leaves elliptic, linear-lanceolate or oblan-
ing a groove from base to apex 10 ceolate, 48(10)mm wide, acute or obtuse 4
9b. Midvein (midrib) on adaxial side of leaves 4a. Adult leaves oblanceolate (widest above the
prominently raised, sometimes fading towards middle) with an obtuse apex. Outer bud scales
apex, but not grooved 11 free but erect, the apices coming together
10a. Adult leaves elliptic-linear (widest at the mid- P. aracensis
dle). Bud scales all imbricate P. hispaniolensis 4b. Adult leaves elliptic to linear-lanceolate (widest
10b. Adult leaves lanceolate to linear-lanceolate at or below the middle) with an acute apex.
(widest below the middle). Bud scales imbri- Outer bud scales spreading P. rusbyi
cate but some outer bud scales may be 5a. Adult leaves (7)813cm long, with an obtusely
spreading P. oleifolius raised, not grooved midrib on adaxial (upper)
11a. Midrib on adaxial (upper) side of leaves slightly side. Receptacles when ripe 812 mm long,
raised, fading towards the acute to apiculate, dark purple P. salicifolius
sometimes pungent apex P. purdieanus 5b. Adult leaves (2)49(11) cm long, with a
11b. Midrib on adaxial side of leaves prominently slightly raised and grooved midrib on adaxial
raised, continuous to the acute or obtuse side. Receptacles when ripe 68mm long, red
(or weakly apiculate) apex 12 or purple 6
12a. Bud scales all imbricate. Adult leaves 410cm 6a. Buds 34mm long, with a few elongated outer
long, lanceolate to linear-lanceolate (widest scales to 10 mm long. Leaf apices all acute.
below the middle) P. guatemalensis Seeds including the epimatium without an api-
12b. Bud scales free at apex. Adult leaves (2.5)4 cal crest P. sellowii
6(7)cm long, elliptic to oblanceolate (widest 6b. Buds 515 mm long, with ovate-cuspidate
at or above the middle) P. trinitensis outer scales all apically elongated. Leaf apices
acute or obtuse. Seeds including the epimatium
Key to the species of Podocarpus with a minute apical crest P. steyermarkii
in South America 7a. Adult leaves (1.5)24(5) mm wide; apex
angustate-acute or pungent 8
When using this key in the field only leaves on 7b. Adult leaves (3.5)415(20) mm wide; apex
branches of mature trees, preferably from sun- acute or obtuse, sometimes acuminate 11
exposed foliage, should be taken into account. Male 8a. Bud scales 46mm long, apiculate or lanceo-
(pollen) and female cones and seeds (they occur on late, outer scales erect or slightly spreading,
dioecious trees) are described as mature. always free at apex 9
8b. Bud scales shorter than 4mm, broadly triangu- 16b. Adult leaves lanceolate-linear to linear, some-
lar, imbricate 10 times narrowly oblanceolate (usually widest
9a. Adult leaves 1.53cm long P. nubigenus below the middle), (3)413(20) cm long,
9b. Adult leaves (2.5)48(9)cm long (5)618(20)mm wide 17
P. parlatorei 17a. Adult leaves (12)1520mm wide, highly vari-
10a. Midrib (midvein) on adaxial (upper) side of able in length but up to 20 cm or more long
leaves continuously raised. Small trees to 12m P. magnifolius
tall P. lambertii 17b. Adult leaves (5)614(18) mm wide, some-
10b. Midvein on adaxial side of leaves a continuous what less variable in length and not longer than
groove, not raised above general surface of leaf. 1213cm 18
828 Trees to 20 m tall but may be dwarfed at the 18a. Midrib on adaxial (upper) side of leaves sharply
highest altitudes P. sprucei and continuously raised, not lying in a (shal-
11a. Bud scales acute, acuminate or cuspidate, erect low) groove. Pollen cones 1015 mm long
and free at apex 12 P. guatemalensis
11b. Bud scales broadly triangular, ovate or rounded, 18b. Midrib on adaxial side of leaves obtusely raised,
imbricate or rarely some outer scales free at lying in a groove or becoming grooved towards
apex 14 leaf apex. Pollen cones 2030mm long 19
12a. Adult leaves 4.57.5cm long, 815mm wide. 19a. Adult leaves pendulous. Pollen cones 67mm
Trees to 30m tall P. celatus wide P. pendulifolius
12b. Adult leaves 24cm long, 48mm wide. Shrubs 19b. Adult leaves spreading. Pollen cones 34 mm
or small trees to 10m tall 13 wide 20
13a. Adult leaves ovate to ovate-linear, 78 mm 20a. Adult leaves with a narrow midrib in a medial
wide (widest below the middle), apex acumi- groove, not elevated above general adaxial
nate P. acuminatus (upper) surface of leaf. Bud scales rounded or
13b. Adult leaves oblanceolate to elliptic, 47 mm ovate, all imbricate P. oleifolius
wide (widest at or above the middle); apex 20b. Adult leaves with an obtusely raised or more or
acute or obtuse P. roraimae less flat and dorsally grooved midrib. Bud
14a. Midrib on adaxial (upper) side of leaves ini- scales (broadly) triangular, with some outer
tially raised but fading towards apex. Pollen scales free at apex P. brasiliensis
cones long and very slender, 2550 1.52mm
P. salignus Key to the species of Podocarpus commonly
14b. Midrib (midvein) continuously raised and/or in cultivation
grooved, or a continuous groove not raised
above general surface of leaf. Pollen cones short The species of Podocarpus keyed out here can be
and slender, or longer than 20mm but then at found in cultivation within and without the areas in
least 3mm wide 15 which they also occur naturally. If a species is only
15a. Pollen cones solitary or more often 26 known to have been planted within the area (in the
together, pedunculate, 612 1.52mm. Seeds sense of the keys) in which it also occurs naturally,
including the epimatium 45 mm long, glo- it is here excluded because then it can be found in
bose, without an apical crest. Buds smaller than the key to the species of that area. Species that are
3mm P. transiens only known or assumed to be grown in a few botanic
15b. Pollen cones always solitary, sessile, more than gardens and/or arboreta are also not included here.
10 mm long. Seeds including the epimatium In cultivation means here: planted in an amenity,
611mm long, not globose and with an apical forestry or horticultural context. When using this
crest (sometimes inconspicuous). Buds usually key in the field only leaves on branches of mature
35mm long 16 trees, preferably from sun-exposed foliage, should
16a. Adult leaves oblanceolate or elliptic (widest at be taken into account. Male (pollen) and female
or above the middle), (1)1.52.5(3.5) cm cones and seeds (they occur on dioecious trees) are
long, 3.55mm wide P. tepuiensis described as mature.
1a. Adult leaves 0.33cm long, 1.45mm wide 2 continuous (if indistinct or fading towards
1b. Adult leaves (1.5)2.514(15)cm long, (3)4 apex, leaves wider than 6mm) 9
15(18)mm wide (if shorter than 3cm, usually 9a. Bud scales on leading shoots long acuminate,
wider than 4mm) 6 812mm P. matudae
2a. Always shrubs with multiple stems 3 9b. Bud scales on leading shoots not long acumi-
2b. Always trees with single stems 5 nate, 25 (rarely to 8)mm long 10
3a. Adult leaves acicular, 1.42mm wide, pungent. 10a. Adult leaves (2)35(7) cm long. Bud scales
Bud scales acute with free apices P. acutifolius erect, with free apices close together. Pollen
3b. Adult leaves ovate-linear to linear, or navicu- cones solitary, 825 68mm P. costalis
late (boat-shaped), obtuse or mucronate. Bud 10b. Adult leaves (2)3.512(15) cm long. Bud
scales (broadly) triangular, imbricate 4 scales spreading or recurved, rarely erect. 829
4a. Adult leaves 45mm wide, with a nearly absent Pollen cones solitary or more often 25
(inconspicuous) midrib on adaxial (upper) together, 1540(60?) 24mm 11
side P. lawrencei 11a. Midrib on adaxial (upper) side of leaves indis-
4b. Adult leaves 22.5 mm wide, with a central tinctly raised, leaf apices obtuse or mucronate.
groove on adaxial side P. nivalis Pollen cones solitary or in pairs. Small bracts
5a. Bud scales lanceolate, to 6 mm long, erect or (foliola) basal to receptacles absent
slightly spreading. Midrib on adaxial (upper) P. latifolius
side of leaves raised but fading towards apex. 11b. Midrib on adaxial side of leaves acutely raised.
Pollen cones 24 together, sessile. Receptacles Leaf apices acute, acuminate or obtuse. Pollen
when ripe 78 mm long; seeds including the cones 25 together (sometimes solitary). Small
epimatium 89 67mm P. nubigenus bracts basal to receptacles present 12
5b. Bud scales obtuse or acute, 23 mm long, 12a. Adult leaves from nearly oval (the shortest) to
imbricate or a few with a free apex. Midvein linear (with parallel margins) in longer leaves
(midrib) on adaxial side of leaves a faint groove. P. polystachyus
Pollen cones solitary or sometimes in pairs, 12b. Adult leaves linear-lanceolate or lanceolate
short pedunculate. Receptacles when ripe 4.5 (widest below the middle), the shortest leaves
5.5 mm long; seeds including the epimatium sometimes oblanceolate (widest above the
33.5 22.5mm P. totara middle) 13
6a. Buds very small (< 3 mm), with imbricate 13a. Bud scales caudate, with a narrow, recurved
scales 7 apex. Shrubs or small trees. Small bracts
6b. Buds larger, with outer scales longer than 3mm (foliola) basal to receptacles also recurved
and spreading or recurved (at least free at apex) P. macrophyllus
8 13b. Bud scales lanceolate or triangular, spreading
7a. Adult leaves narrowly linear-lanceolate, with a more or less straight apex. Potentially
(3)47 mm wide. Pollen cones very slender, large trees. Small bracts basal to receptacles not
2550 1.52 mm. Receptacles when ripe recurved 14
56mm long; seeds including epimatium 78 14a. Adult leaves (4)610mm wide. Pollen cones
45mm P. salignus 1525(30) 22.5 mm. Receptacles when
7b. Adult leaves linear-lanceolate, (5)714(17)mm ripe 916mm long; seeds including the epima-
wide. Pollen cones more robust, 1020 23mm. tium ca. 9mm long, green or glaucous
Receptacles when ripe 1525(30) mm long; P. fasciculus
seeds including the epimatium 1520 14b. Adult leaves (5)615(18) mm wide. Pollen
1215mm P. elatus cones 2040(60?) 23.5 mm. Receptacles
8a. Adult leaves (3)45(7)mm wide; midrib on when ripe 712mm long; seeds including the
adaxial (upper) side of leaves fading towards epimatium (8)1015mm long, purplish green
apex P. elongatus or blackish purple 15
8b. Adult leaves (3)415(18) mm wide; midrib
on abaxial side of leaves acutely raised and
15a. Receptacles subtended by two 26 mm long long and 4mm wide, with a prominent distal crest,
bracts (foliola); epimatium around seeds green. Seed proper not observed.
without a crest. New leaves flushing red
P. neriifolius Distribution
15b. Receptacles subtended by two minute, 12mm
long bracts; epimatium around seeds with a Brazil: Amazonas (Serra da Neblina); Venezuela:
crest. New leaves flushing yellowish green Amazonas (Sierra de la Neblina), Bolivar (Chimant,
P. subtropicalis SW Amuri-tepui).
TDWG codes: 82 VEN 84 BZN-AM
Taxonomic notes
Podocarpus affinis Seem., Fl. Vitiensis: 266. 1868.
It is commonly stated that Thomas Kirk discovered Type: Fiji: Western Division, Viti Levu, Voma Peak,
this species in the upper part of Buller Valley, Nelson, B. C. Seemann 574 (holotype K).
New Zealand (near the outflow of Lake Rotaiti),
but his collection from there is only one element Etymology
cited in the protologue, as he also acknowledges
T. F. Cheeseman for further specimens. At Kew (K) The species epithet means allied to; Seemann
is a sheet with a mixture of male and sterile (female?) thought it was allied to Podocarpus elatus.
branchlets collected by Cheeseman at Lake Rotaiti.
Since the plant is dioecious, these constitute more Vernacular names
than one collection and we do not know what Kirk
saw of this (if any, they are likely duplicates). A kuasi (Viti Levu)
Description Conservation
Small to medium size tree to 1520m tall. Bark Doyle (1998) has listed this species as Endangered
not described. Branches short and spreading; foli- (EN) on grounds of its very limited distribution and
age branches towards ends of main branches slen- apparent decline and fragmentation of its habitat. It
der, terete, terminating in small buds with more was listed as Vulnerable (VU) by the same author in
or less erect, free, narrowly triangular, acute scales. the Global Redlist of Conifers (Farjon & Page, 1999)
Leaves on seedlings similar to those on mature trees, under the 1994 A-criterion which estimates decline
2.55cm long, oblanceolate to elliptic or linear-ellip- only. Since these criteria have changed in 2001, a
tic, 69mm wide, widest at or above the middle, reassessment of this species in 2011concluded that it
832 coriaceous, gradually tapering to a petiolate base; does not meet them.
apex obtuse to rounded; margins slightly revolute, IUCN: NT
lustrous green above, dull brownish green below.
Midrib narrow (ca. 0.5mm wide) and acutely raised Uses
on adaxial (upper) side, similar but fading towards
apex on abaxial side. Stomata very small, in numer- The wood of this species, according to Seemann, was
ous irregular and intermittent lines on abaxial side. used for outrigger canoes by the natives; presum-
Pollen cones and seed cones remain unknown. ably it was used for other purposes as well. Depletion
of the resource has greatly diminished this use at
Taxonomic notes present. The species is not known in cultivation.
Vernacular names
Description Ecology
Small trees 815m tall, to 25cm d.b.h. Bark not Podocarpus atjehensis was discovered and described
described. Branches spreading and ascending, form- (as a variety of P. neriifolius) from the summit area of
ing a pyramidal to rounded crown. Foliage branch- a high mountain in northern Sumatera growing in
lets terete, with narrow ridges from decurrent leaf mossy dwarf forest and shrubland. The type collec-
petioles; leading branches terminating in large buds tion was gathered at 2900m a.s.l. on a slope below
614mm long, 58mm wide, with broadly triangu- the summit of Gunung Kemiri, and it may (or may
lar scales, the outer scales with apiculate, recurved not) occur near or on the summit of nearby Gunung
tips, the inner scales shorter and more or less imbri- Leuser, the highest mountain in the area at 3145m.
cate. Leaves of young plants linear, usually deflected, The other locality reported for this taxon is in New
to 18cm long and 9mm wide; leaves of adult Guinea at 1800m a.s.l.
trees linear-lanceolate, petiolate at base, strongly
deflected, 611cm long, 58mm wide, coriaceous, Conservation
gradually narrowing at both ends; apex long acute;
midrib on adaxial (upper) side less than 0.4mm This species is only known from two very disjunct
wide and acutely raised, on abaxial (lower) side ca. locations. This type of disjunction is also reported
1mm wide and raised to a prominent flat ridge; leaf from other groups of organisms. In both locations
colour dark green above, light green below, flushing there is a considerable human footprint on the land-
pink or rosa. Pollen cones axillary, solitary, sessile, scape and although we do not know to what extent
subtended by broadly ovate-triangular bud scales, this affects this species, it is assumed that some
long cylindrical, 2530mm long, 44.5mm wide at decline has occurred. IUCNs Conifer Specialist
Group therefore wish to flag this species as Near cones axillary, solitary or in clusters of 23, sessile
Threatened. or short pedunculate with several basal bract scales,
IUCN: NT cylindrical, elongating to 3550mm, 2.53.5mm
wide; microsporophylls triangular, spreading, with
Uses two pollen sacs. Seed cones axillary, solitary on a
23mm long peduncle; receptacles with 2 bracts
No uses have been recorded of this poorly known (foliola) 12mm long at base, obliquely bilobate,
species. 6mm long, swelling with a truncate distal end,
green turning red and succulent when ripe. Seeds
including the epimatium ovoid, 6 5mm, the dis-
Podocarpus borneensis de Laub., Blumea 30 tal end cristate, green turning purple or dark brown 837
(2): 266. 1985. Type: Malaysia: Sabah, Sandakan when ripe. Seed proper not observed.
District, Bukit Tawai, D. J. de Laubenfels P 702
(holotype L). Distribution
Podocarpus polystachyus R. Br. ex Endl. var. rigidus Borneo: Indonesia: Kalimantan (including Karimata
Wasscher, Blumea 4 (3): 460. 1941. Island); Malaysia: Sabah, Sarawak.
TDWG codes: 42 BOR-KA BOR-SB BOR-SR
Etymology
Ecology
The species epithet refers to the island of Borneo,
where it is endemic. Podocarpus borneensis occurs in mossy forest on
mountain ridges, often in rocky, exposed sites as a
Vernacular names small or stunted tree. Downslope, where the forest
becomes taller and has a closed canopy, scattered
bisit, bubung, buloh (local names in Sarawak) trees may reach into the canopy and attain 20m
or more. It also occurs in keranga forest on nearly
Description white sand and it was once or twice collected from
swamp forest at low altitude (ca. 350m) on peaty
Small to medium sized trees 510(23) m tall, up to soil. Its more common altitudinal range is from
30cm d.b.h. Bark not described. Branches spread- 700m to 2100m a.s.l. On the Meurong Plateau
ing, forming a rounded or domed crown. Foliage (sandstone), it may become the dominant tree in
branchlets terete, more or less grooved, glabrous, some localities, elsewhere it is mixed with vari-
terminating in 410mm long, 23mm wide buds ous conifers (Agathis, Dacrydium, Dacrycarpus,
with erect or slightly spreading, lanceolate to lin- Podocarpus, Phyllocladus) and angiosperm shrubs
ear outer scales. Leaves on juvenile plants short and trees.
petiolate, linear, to 16cm long and to 14mm wide,
straight or slightly curved, gradually narrowed at Conservation
base, acute or obtuse. Leaves of mature trees often
crowded at the distal end of branchlets, spreading IUCN: LC
or ascending, linear-ovate or elliptic, 37(9)cm
long, 813mm wide, straight or slightly curved, Uses
short petiolate at a gradually narrowing base, taper-
ing to an obtuse or rounded (rarely acute) apex, No economic uses have been recorded of this spe-
thick coriaceous. Midrib acutely raised on adaxial cies. Its ecology and commonly low stature make
side, nearly flat on abaxial side, drying to a groove; it an unlikely target of timber logging. Where it is
leaf colour dark green above, pale green below. abundant, like on the Merurong Plateau, its wood
Stomata very small, in numerous irregular lines on is locally used for construction of houses and other
abaxial (under) side on either side of midrib. Pollen carpentry work. It is not known to be in cultivation.
Podocarpus bracteatus Blume, Enum. Pl. Javae 1: orange-red or bright red and succulent when ripe.
88. 1827. Podocarpus neriifolius D. Don var. Seeds at truncate end of the receptacle, enclosed in
bracteatus (Blume) Wasscher, Blumea 4 (3): 449. a smooth epimatium, green turning purplish green
1941. Type: Indonesia: Jawa, [locality not stated], when ripe, ovoid-oblong or ovoid, 1014mm long,
C. L. Blume s.n. (holotype L). ca. 7mm wide. Seed proper ovoid, 710mm long,
slightly flattened.
Etymology
Taxonomic notes
The species epithet describes the two conspicuous
bracts at the base of the receptacle. Podocarpus bracteatus is quite similar to P. neriifolius,
838 of which Wasscher (op. cit.) considered it to be a
Vernacular names mere variety. However, De Laubenfels (1985, 1988)
not only kept it as a distinct species, but classified
ki marak, ki pantjar, ki putri (Jawa); kayu unung it in a different section: Longifoliolatus as opposed
unung (Sumatera) to Foliolatus. Both are named for the foliola, i.e.
the two basal, subulate bracts below the receptacle.
Description De Laubenfels held that these were about 2mm
long in species of sect. Foliolatus, which contains
Trees to 40m tall, to 1m d.b.h., bole straight, in P. neriifolius, and over 3mm in those of sect.
large trees slightly fluted or rarely buttressed at Longifoliolatus. However, in the widespread species
base. Bark smooth, thin, on large boles with narrow, P. neriifolius basal bracts or foliola as long as 5 or
longitudinal flakes or strips, light brown weather- even 6mm occasionally occur, and consequently
ing grey; inner bark pinkish or reddish brown, this is a spurious character. To base divisions in the
fibrous. Branches spreading, forming a rounded genus on this single feature amounts to artificial
or domed crown. Foliage branchlets terete, more classification (see taxonomic notes above under the
or less grooved, glabrous, terminating in 512mm genus).
long buds with spreading, triangular to lanceolate
outer scales. Leaves on juvenile plants short peti- Distribution
olate, linear-lanceolate, 1523cm long, 1520mm
wide. Leaves of mature trees shorter and nar- Indonesia: Flores, Jawa, N Sumatera.
rower, mostly linear-lanceolate or lanceolate, TDWG codes: 42 JAW LSI-LS SUM
(6)812(14)cm long, 914mm wide, straight or
slightly falcate, gradually narrowing to a petiolate Ecology
base, gradually tapering to an acute apex. Midrib
acutely raised on adaxial (upper) side and 0.4mm Podocarpus bracteatus is a scattered canopy tree of
wide, flat or obtusely (sometimes acutely) raised evergreen montane tropical rainforest, most com-
abaxially; leaf colour dark green above, pale green monly found on forested volcanoes of Jawa at alti-
below, new leaves flushing red. Stomata very small, tudes between 1000m and 2600m a.s.l. It has
in numerous irregular lines on abaxial (under) side occasionally been collected from lower altitudes
on either side of midrib. Pollen cones axillary, soli- between 400m and 1000m a.s.l. Its ecology is simi-
tary or in clusters of 23, sessile with several basal, lar to that of P. neriifolius.
large, ovate-triangular, carinate bud scales, cylindri-
cal, elongating to 2550(60)mm, 3.54mm wide; Conservation
microsporophylls spirally arranged, triangular,
spreading, with two globose pollen sacs. Seed cones Logging of larger trees of this species probably has
axillary, solitary on a 1020mm long peduncle; had an impact on its conservation status, but it
receptacles with 2 subulate bracts (foliola) 45mm seems too widespread and still reasonably common
long at base, swelling to become ellipsoid with a at least in western Jawa. If the botanical distinction
truncate distal end, 1014mm long, yellow turning with P.neriifolius is slight and not often made by
f oresters in the field, it will be very difficult to know lustrous green above, dull green below. Midrib on
whether this species is declining or not. It is at pres- adaxial (upper) side narrow (less than 1mm), con-
ent not considered to be threatened with extinction. tinuous or nearly so, obtusely raised or flat and with
IUCN: LC a central groove, on abaxial side wider, ca. 1mm,
obtusely raised or nearly flat but conspicuous to
Uses apex. Stomata small, in numerous intermittent lines
on either side of abaxial midrib. Pollen cones axil-
This species is an important timber tree in Jawa. Its lary, solitary, sessile, elongating to ca. 30mm, 3mm
excellent wood is used for house construction and wide at anthesis; microsporophylls broadly triangu-
carpentry and for making oars, spars and masts of lar, acute, with two globose pollen sacs. Seed cones
sailing vessels. More specialized uses requiring high axillary, solitary on 512mm long, slender pedun- 839
grade timber are veneer, furniture making, cabinet cles terminating in two minute scales (foliola) sub-
making, interior trim, household utensils, and wood tending the eventually 78mm long, 45mm wide,
carving. As far as known it is not in cultivation. succulent red receptacle consisting of an axis with
two unequal bracts. Seeds solitary, including the epi-
matium 78mm long, subglobose or broadly ovoid,
Podocarpus brasiliensis de Laub., Fl. Venezuela 11 with a weakly developed crest. Seed proper more or
(2): 31. 1982. Type: Brazil: Distrito Federal, Brasilia, less ovoid, 45mm long, grooved or pitted (in sicco).
Horto do Guar, E. P. Heringer 8034 (holotype NY).
Distribution
Podocarpus barretoi de Laub. & Silba, Phytologia 68:
65. 1990. Brazil: Bahia, Distrito Federal, Gois, Mato Grosso,
Minas Gerais, Rondonia, Roraima; Venezuela.
Etymology TDWG codes: 82 VEN 84 BZC-DF BZC-GO BZE-BA
BZL-MG BZN-RO BZN-RM
The species epithet means from Brazil, where the
type specimen was collected. Ecology
Plate 35. Podocarpus brassii. 1. Branch with foliage and pollen cones (var. humilis). 2. Branchlet with
leaves and pollen cones (var. brassii). 3. Branch with foliage and seed cones (var. brassii). 4. Branchlet with
leaves and seed cone (var. brassii). 5. Seed cone. 6, 7. Leaves.
icrosporophylls with narrowed 11.5mm long
m frost. It will probably be a slow growing shrub in cul-
apex, bearing two globose pollen sacs. Seed cones tivation, but should be tried.
towards ends of foliage branchlets, axillary on short
peduncles; receptacles subtended by two 34mm Podocarpus buchii Urb., Feddes Repert. Sp. Nov.
long bracts (foliola), 68mm long, distally bilobed Regni Veg. 19 (1621): 298. 1924. Podocarpus
or truncate when swollen and becoming dark pur- angustifolius Griseb. subsp. buchii (Urb.) Staszk.,
ple. Seeds including the covering epimatium ovoid, Fragm. Fl. Geobot. 33: 77. 1988; Podocarpus aris-
810(12) 67mm, with a small distal crest, dark tulatus Parl. var. buchii (Urb.) Silba, J. Int. Conifer
purple, glaucous or pruinose. Seed proper not Preserv. Soc. 7 (1): 31. 2000. Type: Hispaniola:
observed. Haiti, near La Bellefontaine, L. von Buch 2089b
(holotype B, destroyed?). 843
Distribution
Podocarpus buchii Urb. var. latifolius Florin, Ark.
Malaysia: Sabah (Mt. Kinabalu and surrounding Bot. 25-A (5): 3. 1934; Podocarpus angustifolius
mountain ridges). Griseb. subsp. buchii (Urb.) Staszk. var. latifolius
TDWG codes: 42 BOR-SB (Florin) Staszk., Fragm. Fl. Geobot. 33: 77. 1988.
Ecology Etymology
Podocarpus brevifolius is locally common in upper This species was named after the German botanist
montane to subalpine dwarf forest on Mt. Kinabalu Christian Leopold von Buch (17741853).
and other mountain ridges and heights in the vicinity.
It grows most commonly on ultramafic substrate and Vernacular names
also on granite, among boulders or from crevices. It
has been found on ultramafic rock at 13501450m a.s.l. No common names have been recorded for this
near a copper mine and on the Bambangan Ridge at species.
1900m in lower montane forest. Its altitudinal range
on Mt. Kinabalu proper is between 2100 and 3800m Description
a.s.l., where it is a constituent of a dwarfed forest to
10m tall, dominated by the conifers Phyllocladus Small trees 510m tall, or larger (?); trunk d.b.h. to
hypophyllus and Dacrycarpus kinabaluensis and the 30cm (or to 100cm?). Bark smooth, brown. Branches
umbrella-crowned angiosperm tree Leptospermum spreading, forming a dense crown. Foliage branchlets
recurvum (Myrtacea). Other common woody plants terete, finely grooved between leaf bases, terminat-
in this zone are the conifer Dacrydium gibbsiae (on ing in globose buds with imbricate, rounded, keeled
ultramafics) and the angiosperms Eugenia (Syzygium) scales and apiculate, partly free outer scales. Leaves on
kinabaluensis, Rhododendron buxifolium and Schima saplings larger than on mature trees, to 5cm long and
brevifolia. Near the summit of the mountain only 9mm wide; leaves on mature trees crowded, directed
dwarfed, shrub-like plants of P. brevifolius occur in forward at a narrower than 45 angle towards the
granite crevices. distal parts of branchlets but more widely spreading
lower down, 1.54cm long, straight, elliptic to linear-
Conservation oblanceolate, (4.5)69mm wide, gradually tapering
to a near sessile to short petiolate base, abruptly nar-
IUCN: NT rowing to a spinescent apex, the spine 11.5mm long;
margins slightly revolute, texture coriaceous, stiff; leaf
Uses colour dark green above, paler green below. Midrib
inconspicuous on adaxial (upper) side, slightly raised
No uses are recorded of this species. It can be above leaf base but soon becoming a groove towards
assumed that provenances from high altitude (above apex, continuous, obtusely raised and wider (0.7mm)
3000m) may be hardy as they experience occasional on the abaxial side. Stomata very small, in numerous
intermittent lines on either side of abaxial midrib. Uses
Pollen cones not observed. Seed cones axillary, soli-
tary on 68mm long peduncles; receptacles an axis No uses have been recorded of this species. If reports
with two fused, unequal bracts, growing to 810mm about large trees with d.b.h. to 1m are correct, such
long, becoming swollen and succulent, bright red. trees may well have been logged and the wood used for
Seeds solitary, elliptical, including the epimatium general construction, carpentry and furniture making.
78mm long, 44.5mm wide, with a prominent,
recurved distal crest. Seed proper not observed.
Podocarpus capuronii de Laub., Adansonia 11 (4):
Taxonomic notes 713. 1971. Type: Madagascar: Fianarantsoa Prov.,
844 Itremo Massif, Mt. Ambatomenaloha, R. Capuron
The leaves of P. buchii var. latifolius in the type 11566 (holotype P).
specimen at K (E. L. Ekman 4913, isotype K) from
the Massif du Nord in Haiti are 1.54cm long and Podocarpus woltzii Gaussen, Bull. Soc. Hist. Nat.
59mm wide. This is overlapping the range of Toulouse 110: 123. 1974; Podocarpus capuronii de
dimensions found in specimens of P. buchii Florin Laub. var. woltzii (Gaussen) Silba, J. Int. Conifer
ascribed to the species, but reaching 23mm wider. Preserv. Soc. 7 (1): 31. 2000.
This does not seem to warrant taxonomic distinction,
as the other charcters observed in the type specimen Etymology
are similar to the species. Gray & Buchholz (1948)
in their revision of American species of Podocarpus This species has been named after R. Capuron, who
included material with much larger leaves in this collected the type specimen.
variety, which appears to belong to P. aristulatus.
Vernacular names
Distribution
No common names have been recorded for this
West Indies: Hispaniola: Dominican Republic, Haiti. species.
TDWG codes: 81 DOM HAI-HA
Description
Ecology
Shrubs or small trees to 20m tall, usually less than
Podocarpus buchii occurs in the mountain forests 6m. Bark breaking in small irregular flakes, brown.
of Hispaniola. The northern population in Haiti, Foliage branchlets terete, with grooves and fine
described by Florin (op. cit.) as P. buchii var. latifolius, ridges between the leaf bases, terminating in small
occurs on limestone massifs. In most populations in buds with elongated, incurved outer scales. Leaves
the south of this country the geological substrate is crowded towards ends of branchlets, spreading for-
also a hard karst limestone with a lateritic thin soil. ward at ca. 3545 or some lower leaves spreading
In the Dominican Republic the species is known wider, linear, (1.5)2.55(7.5)cm long, 25mm wide,
from two locations in montane forest on metamor- straight or slightly falcate, gradually narrowing to
phic rock types. The forest is mostly evergreen tropi- a sessile or short petiolate base, more abruptly con-
cal rainforest dominated by angiosperms with small verging distally to an obtuse or minutely acuminate
leaves on steep, rocky slopes, ridges and summits at apex; margins mostly parallel, slightly revolute; tex-
altitudes between 600m and 2050m a.s.l. ture coriaceous; leaf colour dark green above, flushing
leaves bright glaucous blue. Midrib thin, raised proxi-
Conservation mally but fading towards apex on adaxial (upper)
side, continuous and markedly visible on abaxial
This species is apparently scattered and is estimated side. Stomata small, in numerous intermittent lines
to have declined by at least 50% while threats to its on either side of abaxial midrib. Pollen cones axillary,
remaining population have not ceased. solitary or with 23 together on 48mm long pedun-
IUCN: EN (A2a, c, d) cles, subtended by scarious bud scales, cylindrical,
elongating to 12(3)cm, 34mm wide at anthesis; and R. Capuron 27065 (type of P. woltzii) in the Paris
microsporophylls 11.5mm long and ca. 1mm wide, Herbarium (P), which were collected in the 1920s
with minutely erose-denticulate margins, each bear- and 1960s. Habitat loss is considered to be very high
ing two subglobose pollen sacs. Seed cones axillary, in the whole of central Madagascar. Deforestation,
solitary, on a short peduncle, receptacle formed of an habitat loss, cutting for firewood, fires, and grazing
axis with two unequal bracts, swelling only slightly to are the main threats to this species. This species is
34mm long and 2.5mm wide. Seeds including the not known to occur in a protected area.
epimatium 1518mm long, 1012mm wide, obliquely IUCN: EN [B2ab (iv)]
ovoid with a distal crest, brown to bluish pruinose or
whitish pruinose. Seed proper not observed. Uses
845
Taxonomic notes No uses have been recorded of this species. It is likely
to have been used as firewood when still abundant;
Podocarpus woltzii was described by Gaussen (op. perhaps the wood of larger trees was used in local
cit.) as a distinct species, but with characters said to construction and hand tool carpentry.
be close to P. rostratus. The main difference stated
seems to be the occurrence inside the leaves of hori-
zontal sclereids, hard bodies that are sometimes Podocarpus celatus de Laub., Fl. Venezuela 11 (2):
interspersed between the mesophyll cells. Sclereids 35. 1982. Type: Bolivia: Potos, Morro, R. W. Pearce
in leaves of conifers occur erratically and are not s.n. (lectotype K, specimens B, designated here).
reliable taxonomic characters, although some spe-
cies may never develop them. Podocarpus woltzii Etymology
was also described with larger leaves than those of
P. rostratus. The longer and wider leaves seem to be The species epithet (Latin: celatus = concealed, hid-
based on material belonging to P. capuronii de Laub. den) relates to the species previously not being rec-
described two years earlier and, like P. woltzii, based ognized as distinct from P. magnifolius.
on a collection made by R. Capuron on the same day
but on a different mountain. Vernacular names
Distribution Description
Bolivia: La Paz, Potos; Brazil: Gois, Mato Grosso; Small trees to 8m tall; trunk to 20cm d.b.h. Bark
Colombia; Ecuador; Peru: Amazonas, Junin, Loreto, smooth, fibrous, brown weathering to grey-brown.
Montaa, Puno; Venezuela: Amazonas, Bolivar, Branches ascending, foliage branches in mature
Tachira. plants dense, forming a (broad) columnar crown;
ultimate branchlets angular, glabrous, terminating in tion. Its altitudinal range is quoted as near sea level
a small conical bud with carinate, somewhat tapered, to 1000m, but this is based on no more than two
to 3mm long and occasionally spreading scales. (or three) localities cited in the literature. This habi-
Leaves densely crowded, spirally arranged, spreading tat note is quoted from S. Y. Hu (1964), but was not
to erect and rigid, sessile to short petiolate, leaf specified with the collected specimens from Jiangsu
blade linear-lanceolate to oblanceolate, 1.23.5cm and Zhejiang (the Sichuan specimen was not seen).
long, 2.54.5mm wide, gradually tapering to base, It is apparently very rare.
in adult leaves with an obtuse apex, midvein acutely
raised adaxially, not raised and wider abaxially; leaf Conservation
colour olive green above (adaxially), glaucous green
below. Stomata conspicuous, in numerous regular Too little is known about this taxon, apart from its 847
lines on abaxial side on either side of the midvein. apparent rarity, to evaluate its conservation status. It
Pollen cones axillary, sessile, in clusters of (1)23 is with certainty known from two localities, one in
per axil, subtended by several scarious, more or less Jiangsu and the other in Zhejiang; a third herbarium
triangular bracts, long cylindrical, 1525mm long, collection was mentioned by S. Y. Hu (op. cit.) to be
1.52mm wide; microsporophylls spirally arranged, from Sichuan. That would be far to the west of the
imbricate, spreading at anthesis, the laminar part other two, neither of which has been specified to
apiculate, 0.50.7mm long, with two basal pollen modern topographic locality names, so they remain
sacs. Seed cones and seeds not known. essentially located as to the provinces only.
IUCN: DD
Taxonomic notes
Uses
This species was originally described as a variety
of the wide-spread species Podocarpus macrophyl- No uses are known of this species; it may be in culti-
lus (Thunb.) Sweet by N. E. Gray (op. cit.), based vation in China locally.
on the collection R. C. Ching 2477 of which she had
seen a specimen in the Herbarium of the Arnold
Arboretum at Cambridge, Massachusetts. S. Y. Hu Podocarpus confertus de Laub., Blumea 30 (2):
(op. cit.) described it as a new species, based on the 271. 1985. Type: Malaysia: Sabah, Tawau District,
same type collection (R. C. Ching 2477) so in fact Gunung Silam, D. J. de Laubenfels P 691
elevating Grays variety in rank to a species. Two (holotype L).
more collections were cited by Hu (Gray only knew
the type), C. L. Tso 1477 from Jiangsu and W. P. Fang Etymology
14261 from Sichuan. A duplicate of C. L. Tso 1477 is in
the Kew Herbarium (K); the collection from Sichuan The species epithet means pressed close together,
could not be verified. This taxon was regarded as a but what is considered to be so is not clear from the
cultivar of Podocarpus macrophyllus (cv. Chingii) by original description or the specimens seen.
S. Y. Zhang in Flora of Zhejiang Province 1 (Lin &
Zhang, 1993), but there is no evidence of it having Vernacular names
been found (only) in cultivation.
No common names have been recorded for this
Distribution species.
Philippines: NE Luzon coast?, Bucas Islands off No common names have been recorded for this
NE Mindanao to Batan Islands in the Luzon Strait; species.
Taiwan: Lanyu (Orchid Island).
TDWG codes: 38 TAI 42 PHI Description
Plate 36. Podocarpus elatus. 1. Habit of tree. 2. Branchlet with leaves and pollen cones. 3. Pollen cones.
4. Branchlet with leaves and seed cones. 5. Seed cone with two seeds.
obtuse on adaxial (upper) side, varying in width and has a warm climate; it becomes rare in drier
with leaf but up to 1mm wide, more flattened on forest in the north of Queensland. These forests
abaxial side; leaf colour lustrous dark green above, are regionally known as complex mesophyll vine
pale green below. Stomata numerous, small, in two forests, characterized by trees with medium-size,
bands of intermittent lines on abaxial side. Pollen entire margined leaves, an abundance of climb-
cones clustered, axillary with 24 together, sessile, ing plants (lianas), and locally palms. The altitu-
cylindrical, 1020mm long, 23mm wide; micro- dinal range of this species is from near sea level
sporophylls very small, imbricate, broadly triangu- to ca. 1000m a.s.l. The soils are usually deep,
lar, with two basal globose pollen sacs. Seed cones alluvial sands and silts and this species is often
axillary, solitary on stout, 310mm long peduncles; found on river banks. Associated species are e.g.
860 receptacles subtended by two very small bracts Castanospermum australe, Grevillea robusta, and
(foliola), growing and swelling to 1525(30)mm nearer the coast in a type called mixed notophyll
long and 1520mm thick at the distal end, ripen- vine forest Alphitonia excelsa, Cryptocarya trip-
ing from red to pruinose dark purple or blue-black, linervis, Cupaniopsis anacardioides, Diploglottis
very succulent. Seeds 1(2) on distal part of recep- cunninghamii, and the palm Livistona australis,
tacle, obliquely attached, ovoid-globose with a short the latter often along streams.
proximal beak and an obscure distal crest, 1520
1215mm when mature; epimatium ripening from Conservation
dark glaucous green to purplish black with a whitish
bloom. Seed proper not observed. IUCN: LC
Ken Hill in Flora of Australia 48: 556 (1998) indicated The wood of this species is resistant to termites and
a collection made by Robert Brown with No. 3117, marine borers like barnacles, it is highly durable
with duplicates at BM and K, as syntype. Endlicher and strong. It is therefore valued and used for con-
(1847) merely referred to Brown as the author of the struction like jetties and boat pilings in salt water,
species and Habitat in Nova Hollandia orientali but for boat construction, packing cases, carpentry,
not to his collections of specimens. We do not know joinery, and wood turning, and also for furniture,
what Endlicher, who was based in Vienna, has seen, especially outdoor furniture. It takes a fine polish
as the conifer specimens at W were all destroyed in and is therefore used in high quality products that
World War II. Assuming Brown 3117 is a collection last long. This species is occasionally planted as a
of which Endlicher saw a duplicate at W and that park or street ornamental tree; in deforested areas
he may have seen other material collected by Brown turned over to agriculture mature individuals may
along the Hunter River as well, the BM duplicate is have been retained as amenity trees. It is obviously
here designated as the lectotype of Podocarpus elatus only suitable for these purposes in warm temperate
R. Br. ex Endl., with an isolectotype at K. to subtropical climates.
Distribution
Podocarpus elongatus (Aiton) LHerit. ex Pers.,
Australia: New South Wales, Northern Territory, Syn. Pl. 2 (2): 580. 1807. Taxus elongata Aiton,
Queensland. Hort. Kew. 3: 415. 1789. Type: South Africa: Cape
TDWG codes: 50 NSW-NS NTA QLD-QU Province, [Caput Bona Spei], F. Masson s.n.
(lectotype BM).
Ecology
Etymology
Podocarpus elatus occurs naturally in coastal and
subcoastal rainforest in the region of the east- The species epithet means elongated and refers to
ern Australian coast which receives ample rain the leaf shape.
Vernacular names low longitudinal grooves was reported by Leistner
in Flora of Southern Africa 1: 40 (1966) and repeated
Breede River yellowwood; Brerivier geelhout in several compilations on southern African trees.
(Afrikaans) My observations of herbarium specimens at K with
a powerful Leica MZ6 binocular microscope at
Description 3040 magnification did reveal a few short lines
of stomata on that side on only some specimens (e.g.
Shrubs or small trees usually 36m tall, but on occa- Pearson 5328 from near Oliphants River). Nearly all
sion attaining 20m, d.b.h. to 50cm; bark on trees stomata are confined to the abaxial side and most
thin, exfoliating in narrow strips, light brown weath- leaves observed did not have any stomata on the
ering grey. Shrubs spreading wide to 1012m, multi- adaxial surface of the leaves. 861
stemmed, regenerating from fires. Foliage branchlets
assurgent, slender, terete, with fine grooves, ter- Distribution
minating in buds 23mm diam. with oblong, nar-
rowly triangular, 46mm long outer bud scales with South Africa: Eastern, Northern and Western Cape
spreading tips. Leaves crowded in upper parts of Provinces; Malawi; Zambia; Zimbabwe.
branchlets, spreading to nearly erect, (1.5)2.55( TDWG codes: 26 MLW ZAM ZIM 27 CPP-EC CPP-NC
7)cm long, (3)45(7)mm wide (to 12cm long and CPP-WC
10mm wide on juvenile plants or vigorous epicormic
shoots), narrowly oblong-elliptic to linear, gradually Ecology
tapering to a petiolate base, more abruptly to an acute
or sometimes obtuse apex; margins flat to slightly Podocarpus elongatus is an uncommon species
revolute; leaf colour glaucous to greyish green above, growing in woodlands in moist sites, usually along
dull whitish green below; midrib on adaxial (upper) (intermittent) streams and in ravines, or on rocky
side thin, raised in lower half and fading towards sites with sparse vegetation. Woodland species in
apex, on abaxial side continuous and distinctly raised. these ravines are Cunonia capensis and Olea capen-
Stomata in two bands of many white lines separated sis, among other trees, and sclerophyllous shrubs
by midrib on abaxial side, only occasionally a few on the drier edges. In the Western Cape its habi-
short lines of stomata on adaxial side. Pollen cones tat contacts with fire-prone vegetation types such
axillary, solitary or sometimes in clusters of 25, ses- as fynbos and as a consequence individual trees
sile or sometimes pedunculate, cylindrical, 1525mm are frequently burnt. Resprouting from the base,
long, elongating to 3040mm at anthesis, 35mm they then develop into broad spreading shrubs or
diam., subtended by ovate or broadly rounded, scari- bushes, while only individuals that are protected
ous, acuminate bud scales; microsporophylls spirally from fire, e.g. by growing in a deep ravine, can
insertend, imbricate, outer part triangular, minutely develop into monopodial trees of some size. In the
denticulate or lacerate, each bearing two oblong basal NE part of its scattered range it is usually a com-
pollen sacs. Seed cones axillary, solitary, on 412mm ponent of moist evergreen forest and grows more
long, stout (1mm diam.) peduncles; receptacles an often into a tree.
axis with two fused bracts, green but swelling to a
scarlet, 915mm long and 1015mm wide (at distal Conservation
end) succulent body. Seeds including the epimatium
712mm long, ellipsoid to ovoid with a slightly nar- IUCN: LC
rowed distal end, glaucous green. Seed proper ovoid
with purple-mottled seed coat. Uses
Taxonomic notes Due to its small size and usually bushy habit, this
species is not of economic importance as a timber
The presence of stomata on the adaxial (upper) side tree. It is rare in cultivation, but a few cultivars have
of the leaves in 1 to several short rows...in shal- been recorded.
Podocarpus fasciculus de Laub., Blumea 30 (2): to purple-green. Seed proper ca. 7 5mm, slightly
277. 1985. Type: Taiwan: Taichung Co., Tashue flattened, brown.
Shan, [Tai-shu Shan], D. J. de Laubenfels P 675
(holotype L). Taxonomic notes
Podocarpus macrophyllus (Thunb.) Sweet var. liuki- De Laubenfels (op. cit.) described P. fasciculus as
uensis Warb., Monsunia 1: 192. 1900; Podocarpus a new species based on his own collections from
macrophyllus (Thunb.) Sweet f. grandifolius Pilg., in Taiwan, with one of these designated as the type.
Engler, Pflanzenr. IV.5 [18]: 80. 1903. Similar plants were described from southern Japan
(Kyushu: Nagasaki; Ryukyu Islands: Iriomote
862 Etymology Island) as P. macrophyllus var. liukiuensis Warb. from
Kyushu, and as P. macrophyllus f. grandifolius Pilg.
The species epithet (Latin fasciculus = cluster or fas- from the Ryukyu Islands. Iriomote Island is close to
cicle) refers to the clustered pollen cones. Taiwan and apparently P. fasciculus occurs on sev-
eral other islands in the Ryukyus, up to the main
Vernacular names islands of Japan.
Etymology Ecology
The species epithet commemorates Lilian Suzette Podocarpus gibbsiae occurs on mountain ridges
Gibbs (18701925), who collected and studied the at altitudes between 1200m and 2400m a.s.l. in
flora of Mt. Kinabalu. mossy forest within the cloud belt on Mt. Kinabalu.
It appears to be confined to ultramafic soil derived
Vernacular names from serpentine and similar rocks. This forest type
has an open canopy up to 2025m tall and consists
No common names are recorded for this species. of a mixture of angiosperms and gymnosperms
(mostly conifers); common conifers are Phyllocladus
Description hypophyllus and Dacrydium gibbsiae. Epiphytes,
from lichens, mosses, and ferns to orchids are
Small to medium sized trees 720m tall. Bark numerous.
becoming scaly on largest stems, brown weather-
ing grey. Crown in more or less sheltered trees coni- Conservation
cal, becoming rounded with age, otherwise usually
irregular. Foliage branches spreading to assurgent, The near endemic population known on
densely leaved distally, forming tufts of foliage; Mt.Kinabalu of this species occupies an area (AOO)
shaded branches with more widely spaced leaves. of less than 100 km2 and the trees are restricted to
Terminal buds on leading shoots 49mm long, a specific soil type, which is discontinuous within
23mm wide, narrowly conical, with free spread- that area. Most trees are now within Mt. Kinabalu
ing, acuminate outer scales. Leaves on young plants National Park. Although occurring within the cloud
36(9)cm long, 47(9)mm wide, linear-lanceo- belt, exceptional spells of drought and the greatly
increased tourism on the mountain pose a potential cones axillary, solitary, cylindrical, 1520mm long,
fire hazard, to which the species is not adapted. 22.5mm wide, with a few rounded scales (from
IUCN: VU (D2) the buds) on a 2mm long peduncle; microsporo-
phylls rounded at apex, with two globose pollen
Uses sacs at base. Seed cones axillary, solitary; recep-
tacles with two 1.5mm long basal bracts (foliola),
No uses have been recorded for this tree. 68mm long and swollen when ripe, becoming pur-
ple. Seeds including the epimatium ovoid-globose,
67 5mm, with a crest at maturity, glaucous green
Podocarpus glaucus Foxw., Philipp. J. Sci. 2: 258. to purplish green; seed proper not observed.
864 1907. Type: Philippines: Mindoro, Parabatugan,
Mt. Halcon, E. D. Merrill 5672 [holotype PNH Distribution
(destroyed?), isotypes K, NY].
Malesia: Admiralty Islands (Manus), Maluku
Etymology [Moluccas] (Seram), Philippines (Mindoro),
Sulawesi; Papuasia: Bismarck Archipelago, New
The species epithet refers to the bluish (glaucous) Guinea, Solomon Islands.
leaves of the type specimen; not all leaves of all TDWG codes: 42 MOL PHI SUL 43 BIS NWG-IJ NWG-
plants of this species are so coloured. PN SOL-SO
The species epithet refers to the clustered pol- Podocarpus glomeratus is a species of the high Andes
len cones, collected closely together into a head occurring in high montane to subalpine forests and
(Stearn, 1983). woodland or scrub, at altitudes from 1800m a.s.l.
(but usually not below 2500m) to 3600m a.s.l.
Vernacular names or perhaps higher. Above 3000m this species is
dwarfed and shrubby; below this it is a constituent
Intimpa, Huampo (Quechua); pino de monte of cloud forest rich in epiphytes, especially mosses
(Spanish) and lichens.
866
Description Conservation
Distribution Description
palo de cruz, palo de puntella (Spanish) This species is only known from five locations, all
visited for collection of herbarium specimens in the
Description 1980s. Its total area of occupancy (AOO) is estimated
to be less than 500 km2 and due to deforestation sev-
Trees 1520m tall; trunk massive, d.b.h. 11.5m in eral populations are fragmented. Logging of large
large specimens. Branches spreading and drooping, trees has had an impact on the number of mature
forming a wide, domed crown. Bark not described. trees, which have become scarce in some areas.
Foliage branchlets slender, terete, with fine grooves IUCN: EN [B1ab (iv)]
Uses wide; microsporophylls with a triangular, ca. 1mm
long apex and two basal, globose pollen sacs. Seed
This species is being logged together with other large cones axillary on a short, 23mm long peduncle;
trees in the forests where it occurs. The wood is used receptacles formed of an axis with two unequal
for general construction and carpentry. It is not bracts and enlarging to only 35 2.54mm, becom-
known to be in cultivation. ing coriaceous. Seeds solitary, including the thick
epimatium 1115mm long, 912mm wide, ovoid-
globose, with a small, inconspicuous crest below the
Podocarpus humbertii de Laub., Adansonia 11 (4): apparent apex.
714. 1971. Type: Madagascar: Antsiranana Prov.,
872 Anjanaharibe Massif, Mont Anjanaharibe, slopes Distribution
and north summit, west of Andapa, H. Humbert et
al. 24741 (holotype P). Madagascar: Antsiranana Province.
TDWG codes: 29 MDG
Etymology
Ecology
This species has been named after H. Humbert, who
collected it in 1951. Podocarpus humbertii is reported from sub-humid
forest, dry lowland deciduous forest, and ericoid
Vernacular names thickets or wooded heath on mountain summits of
gneiss and granite. The elevation ranges from 1600m
No common names have been recorded for this to 2410m a.s.l. according to data on herbarium speci-
species. men labels. In the original description of the species it
was reported to occur up to 2800m a.s.l.
Description
Conservation
Small, shrubby trees 315m tall or perhaps larger.
Bark exfoliating in small flakes, brown weathering This species is know from five to seven subpopula-
grey. Foliage branchlets spreading to erect, slender, tions (locations) and most of the herbarium collec-
terete, with persisting leaf bases, terminating in tions were made between 19501960 with only one
small, subglobose buds with imbricate, triangular, recent collection in 2001. Based on the mapped her-
obtuse scales. Leaves small, crowded towards ends barium collections, the area of occupancy (AOO)
of branches, spreading at 50 or less from shoot, is calculated as <150 km2, which falls within the
curved downward towards apex, overlapping each threshold of Endangered (EN). Its habitat is (in part)
other particularly near ends of branchlets and con- dry lowland deciduous forest, which is under pres-
cealing buds, obovate to oblanceolate (the wid- sure from grazing. A continuing decline is inferred
est part towards the apex), 715mm long, 24mm from the collection dates and the degradation of
wide, gradually narrowing towards a petiolate base; parts of its habitat. This species has been collected in
margins flat or slightly revolute, abrubtly narrowing the following protected areas: Tsaratanana Reserve,
to form an obtuse, rounded or sometimes acumi- Marojejy National Park, and Anjanaharibe-Sud
nate apex; texture coriaceous; leaf colour dark green Special Reserve. Only one collection, from Mont
above, dull green below. Midrib narrow, inconspic- Tsaratanana, was made recently, in 2001.
uous and discontinuous or in a groove on adaxial IUCN: EN [B2ab (iii, iv)]
(upper) side, continuous and wider but flattened
on abaxial side. Stomata small, in numerous inter- Uses
mittent white lines on either side of abaxial midrib.
Pollen cones axillary, solitary or in pairs on a 36mm No economic uses have been recorded of this spe-
long peduncle, cylindrical, 820mm long, 2.53mm cies. It is probably used for firewood locally.
Podocarpus insularis de Laub., Blumea 30 (2): globose pollen sacs. Seed cones axillary, solitary on
266. 1985. Type: Papua New Guinea: Louisiades a 510mm long peduncle; receptacles a short axis
Archipelago, Tagula Island [Sudest Island], to 10mm long with two fused bracts, swelling to
L. J. Brass 27987 (holotype L). become red and succulent when ripe. Seeds solitary
at truncate end of receptacle, enclosed in a smooth
Etymology epimatium, green turning purplish when ripe, ovoid
to subglobose, 1015mm long, without a crest. Seed
The species epithet refers to its insular (on islands) proper not observed.
distribution.
Taxonomic notes
Vernacular names 873
This species appears to be more variable in its leaf
In Papua New Guinea this species is known locally shapes and size than described by De Laubenfels
as dala and tunum (Milne Bay); in the Solomon (1985, 1988), at least in material seen at the Kew
Islands dengali or dingale and mou were recorded Herbarium (K) that seems to be best assigned to it.
by collectors. Unfortunately, most specimens are sterile, and both
pollen cones and seed cones have been described
Description here more or less provisionally.
Plate 37. Podocarpus lambertii. 1. Habit of tree. 2. Branchlet with leaves and pollen cones. 3. Pollen
cones. 4. Branch with leaves and seed cones. 5. Seed cone. 6. Leaf, upperside. 7. Leaf, underside.
Conservation in stunted trees. Foliage branchlets stout, terete or
slightly angular, finely grooved from the decurrent
This species has a large range; the extent of occur- leaf bases, terminating in obtuse buds 23.5mm
rence (EOO) calculated from mapped herbarium diam. with narrowly triangular, acuminate, some-
specimens = 761,575 km2. The human footprint fac- times recurved outer scales. Leaves on juvenile
tor is fairly high (45), especially near the coast, where plants up to 17cm long and 20mm wide. Leaves on
habitat and individual trees are likely to have disap- mature trees shorter but variable and often crowded
peared. The IUCN Conifer Specialist Group feels it towards ends of branchlets, (2)3.58(12)cm long,
is warranted to flag this species as NT. The species is mostly straight, (5)612(18)mm wide, elliptic to
not known to occur in any protected area. linear-elliptic or linear, longer leaves with parallel
876 IUCN: NT sides in middle part, gradually or more abruptly
narrowing to a petiolate base, abruptly narrowing to
Uses an obtuse or mucronate to apiculate apex; margins
slightly revolute; leaf colour dark green to glaucous
No commercial uses have been recorded for this green above, dull green below; flushing leaves bright
small tree, of which the timber has not much value. green or bronze. Midrib thin and inconspicuous on
It may be used locally for fence posts and most likely adaxial (upper) side, fading towards upper part,
for firewood. It is not known to be in cultivation. more prominent and continuous on abaxial side.
Stomata in numerous intermittent lines on either
side of abaxial midrib, rarely a few present on upper
Podocarpus latifolius (Thunb.) R. Br. ex Mirb., leaf surface. Pollen cones axillary, solitary or rarely
Mm. Mus. Hist. Nat. 13: 75. 1825. Taxus lati- in pairs, sessile or on very short peduncles, cylin-
folia Thunb., Prodr. Pl. Cap.: 117. 1800. Type: drical, elongating to 2030mm long at anthesis,
South Africa: Cape Province, [Houtniquas, 34mm wide, subtended by 34mm long, narrowly
Grootvadersbosch, aliis], C. P. Thunberg UPS triangular, keeled bud scales; microsporophylls
23780 (holotype UPS). Fig. 270 triangular to broadly ovate, lacerate, bearing two
oblong pollen sacs. Seed cones axillary, solitary on
Podocarpus latifolius (Thunb.) R. Br. ex Mirb. var. slender 515mm long peduncles; receptacles com-
latior Pilg., in Engler, Pflanzenr. IV.5 [18]: 90. 1903; posed of an axis with two bracts, one or sometimes
Podocarpus latior (Pilg.) Gaussen, Trav. Lab. Forest. both fertile, swelling to 814mm long and 812mm
Toulouse T. 2, 1 (2, 21): 66. 1976 (nom. inval., Art. 33.2). wide and becoming succulent, turning from glau-
cous green to pink and sometimes reddish or bluish
Etymology purple when ripe. Seeds including the epimatium
obovoid to subglobose, 711mm long, slightly apic-
The species epithet latifolius means with broad ulate or crested, glaucous green turning purple or
leaves. violet. Seed proper smooth, the seed coat contain-
ing resin cavities.
Vernacular names
Distribution
Broad-leaved Yellowwood, True Yellowwood;
Opregte geelhout (Afrikaans); umGeya (Xhosa) South Africa: from the Cape to the Limpopo
Province.
Description TDWG codes: 27 CPP-EC CPP-WC LES NAT OFS
SWZ TVL-GA TVL-MP TVL-NP TVL-NW
Trees to 30m tall or more in forests, or stunted small
trees 23m tall on exposed mountain slopes, bole of Ecology
large trees to 1.2m d.b.h. Bark thin, smooth, light
brown, exfoliating in long strips, weathering grey. Podocarpus latifolius is a canopy forest tree in the
Crown relatively small in forest trees, low spreading coastal and midland primary forests where there is
sufficient rainfall and natural protection from fires Vernacular names
to allow such forest types to develop. In open coastal
bushland and on dry, rocky mountain slopes it only No common names have been recorded for this
grows to a stunted tree a few meters tall at most. It species.
occurs from near sea level to 2000m a.s.l. In for-
ested valleys near the coast it can be associated with Description
Afrocarpus falcatus and both are there commonly
emergents above a lower canopy of angiosperm trees, Trees to 40m tall, bole erect, straight, to 80cm d.b.h.
among which members of the families Celastraceae, Bark nearly smooth, soft, exfoliating in thin strips,
Araliaceae and Flacourtiaceae, as well as Olea capen- light brown. Branches spreading, in mature trees
sis are often seen. forming a rounded crown. Foliage branchlets terete, 877
glabrous, finely grooved or striated, terminating an a
Conservation small, globose bud up to 4 4mm, with triangular,
imbricate scales forming a short conical apex, the tips
IUCN: LC of the scales at apex often recurved. Leaves on saplings
and young trees usually larger than on mature trees;
Uses shaded leaves also larger than sun-exposed leaves on
mature trees, to 25cm long and 25mm wide. Leaves
Broad-leaved Yellowwood is a valuable timber on mature trees exposed to sun (7)1019cm long,
tree producing even-grained, light weight, pale 1020(25)mm wide, long petiolate, linear-lanceo-
yellow wood suitable for a variety of purposes. It late to linear, straight or slightly curved, more or less
was extensively used in colonial times for railway abruptly widening above the 615mm long petiole,
sleepers and construction, and many houses were more gradually tapering to an acute or more com-
built with it, such as the old Cape homesteads of monly acuminate apex. Midrib on adaxial (upper)
which many still exist. It is still valued for indoor side obtusely raised, ca. 1mm wide, on abaxial side
carpentry and floors, as in former times, but large wider and flatter, fading towards apex. Stomata very
trees have become much less common and smaller small and indistinct, in numerous irregular lines on
sizes are now used for furniture making, especially abaxial side. Pollen cones axillary, clustered with 35
when well seasoned and nicely figured with darker on a short peduncle or sessile, raising from globular
streaks. Africans value the wood for coffins. In buds with rounded scales, cylindrical, elongating to
South Africa this species is commonly planted as an 2040mm or longer, 2,53.5mm wide; microsporo-
amenity tree in parks and along streets. Elsewhere phylls with an elongated apex, bearing two elongated
it is uncommon and mainly represented by speci- pollen sacs. Seed cones axillary, solitary on slender
mens in botanic gardens. It is too slow growing for peduncles (3)1015mm long; receptacles with 2 very
profitable forestry plantation, which is unfortunate short (12mm) bracts (foliola) at base, ca. 10mm
as that role is mostly taken by several species of long. Seeds including the epimatium at least 10mm
Eucalyptus, some of which have turned out to be long but fully mature phase not observed.
invasive pests.
Distribution
Podocarpus laubenfelsii Tiong, Blumea 29 (2): 523. Borneo: Malaysia: Sabah (Mt. Kinabalu, Mt.
1984. Type: Malaysia: Sabah, Ranau District, Trusmadi), Sarawak (Lawas); Indonesia: Kalimantan
Mt. Kinabalu N. P., Bukit Burong, D. J. de Timur (Gunung Palimasan).
Laubenfels P 715 (holotype L). Fig. 269 TDWG codes: 42 BOR-KA BOR-SB BOR-SR
Etymology Ecology
This species has been named in honour of David J. Podocarpus laubenfelsii occurs scattered in keranga
de Laubenfels, a long-time student of Podocarpaceae. forest with Agathis borneensis, Nageia w allichiana,
Sundacarpus amarus, Dacrydium gracile, and Etymology
Falcatifolium falciforme, often on nutrient-poor and/
or water-logged, acidic soils. The species is also scat- This species was named after Robert William
tered in primary rainforest and mossy forest; grow- Lawrence (18071833), who collected plants in
ing as a large emergent tree on rocky ridges; it may be Tasmania around 1826.
more common in heath forests at higher elevations.
Altitude of P. laubenfelsii ranges from 920m to 1650m Vernacular names
a.s.l. This species occurs with scattered individuals in
the forest, but can be dominant in heath forests at Mountain plum pine, Plum pine
higher altitudes.
878 Description
Conservation
Procumbent or more or less erect shrubs to 4m
Most collections of this species are from Mt Kinabalu tall, usually creeping and prostrate, forming 45m
National Park. Deforestation is widespread outside of wide clumps. Branching often profuse, with densely
protected areas in the region, indicating a consider- set foliage branchlets and in creeping shrubs with
able decline in the past that has not abated. Outside long leaders (whip shoots), terminating in small,
Mt. Kinabalu National Park it is known from only globose buds 12mm diam. with broadly trian-
three locations, one just outside the park, the other gular, imbricate scales. Leaves sessile, decurrent,
two at considerable distance. Its total range is probably 416mm long, linear-oblong, the shortest leaves
incompletely known as the species was only described nearly oval, 45mm wide, gradually tapering to a
and named in 1984. It is a large forest tree with consid- slightly twisted, narrow base; apex obtuse, some-
erable timber value. Its only protected location so far times minutely apiculate; midrib nearly absent
known is in Mt. Kinabalu National Park. on adaxial (upper) side, prominent but obtuse
IUCN: EN [B2ab (ii, iii, v)] on abaxial side; leaf colour mid-green or dark
green above, light green below with two grey-
Uses green stomatal bands. Stomata small, in intermit-
tent, wavy lines on either side of abaxial midrib.
This species attains large sizes in primary lower Pollen cones axillary and solitary or in groups of
montane rainforest and is consequently a valuable 23(6) at distal end of foliage branchlets, sessile
timber tree logged and traded as other podocarp or on short peduncles, cylindrical, 47mm long,
trees, without distinction to species or even genus. 22.5mm wide; microsporophylls imbricate, very
Its excellent wood is used for house construction and small, broadly triangular, enclosing towards their
carpentry and for making oars, spars and masts of base two relatively large, globose pollen sacs. Seed
sailing vessels. More specialized uses requiring high cones axillary, solitary, sessile or shortly pedun-
grade timber are veneer, furniture making, cabinet culate; receptacles while growing ca. 3mm long,
making, interior trim, household utensils, and wood swelling strongly at maturity to 56mm, becom-
carving. It is not known to be in cultivation. ing subglobose, succulent and dark red. Seeds
solitary at distal end of receptacle, narrowly ovoid,
4.55mm long, 2.53mm wide, with a narrowing
Podocarpus lawrencei Hook. f., London J. Bot. distal end; epimatium olive green. Seed proper not
4: 151. 1845., [lawrencii] Podocarpus alpinus observed.
R. Br. ex Hook. f. var. lawrencei (Hook. f.) Hook.
f., Fl. Tasmania 1 (5): 356. 1857. Type: Australia: Distribution
Tasmania, R. W. Lawrence 218 (holotype K). Fig. 271
Australia: New South Wales, A. C. T., Victoria,
Podocarpus alpinus R. Br. ex Hook. f., London J. Bot. Tasmania.
4: 150. 1845. TDWG codes: 50 NSW-NS TAS VIC
Ecology Etymology
Podocarpus lawrencei is a subalpine to alpine shrub This species was named after the German botanist
limited to the highest mountains in the southern part Carl Ludwig Ledermann (18751958), who first col-
of the Great Dividing Range and in Tasmania. Its lected it in the Sepik River delta.
altitudinal range is between 1100m and 2030m a.s.l.
and it grows mostly in rocky terrain, e.g. scree slopes, Vernacular names
broken rocky plateaus and ridges formed by acidic
igneous or metamorphic rock types. In the Great sua (Papua); babako (Papua New Guinea mainland);
Dividing Range it may also occur in wet sclerophyll neleel, nelil (New Britain)
forest with Acacia spp., Eucalyptus spp., and Telopea 879
sp., where it can reach to 4m tall; above the tree line it Description
is found in subalpine/alpine dwarf scrub mixed with
alpine herbaceous grassland, some associated species Trees to 25(30) m tall; trunk to 60cm diam., erect.
there are Prostanthera cuneata, Grevillea australis and Bark more or less fibrous, dark brown. Branches
Eucalyptus australis. In Tasmania, three endemic coni- spreading, forming a rounded crown. Foliage
fers are often associated with Podocarpus lawrencei: branchlets terete, glabrous, finely grooved or stri-
Diselma archeri (Cupressaceae), Microcachrys tetrag- ate, terminating in an elongated bud 612mm long,
ona, and Pherosphaera hookeriana (Podocarpaceae). 34mm wide at base, with free, erect to slightly
The angiosperm flora is also distinct, with e.g. Orites spreading lanceolate scales drawn out in a cau-
revoluta, Richea scoparia, Epacris serpyllifolia, and date apex. Leaves lanceolate to linear-lanceolate,
Leptospermum ruprestre. 820(22)cm long, (11)1525mm wide, straight
or slightly curved, distinctly petiolate, more or less
Conservation abrubtly widening at base, with parallel sides for
most of their length, gradually tapering to an acu-
IUCN: LC minate or sometimes acute apex. Midrib distinctly
raised and narrow on adaxial (upper) side, wider
Uses and flattened on abaxial side; leaf colour lustrous
green on both sides. Stomata very small, in numer-
Mountain plum pine is suitable as a low, hardy shrub ous irregular lines on abaxial side on either side of
in countries with mildly cold winters. It is planted in midrib. Pollen cones axillary, on a 34mm long
rockeries or as undergrowth in park-like tree plant- peduncle, in groups of 13, subtended by 45mm
ings and provides an evergreen, spreading shrub long acuminate bracts (bud scales), elongating to
with attractive, red fruits (the ripe receptacles) 3.54.5cm, 4mm wide at anthesis; microsporo-
topped by a shiny, green seed. It is uncommon in phylls apiculate, bearing two elongated pollen sacs
Europe, North America and Japan probably because at base. Seed cones axillary, solitary, on a 515mm
it is not easy to germinate; in Australia and New long peduncle; receptacles subtended by 2
Zealand gardeners seem to have a preference for recurved, 2mm long bracts (foliola) and with one
conifers from the northern hemisphere, too. apical bract visible when still growing, at maturity
1016mm long, ripening to a swollen, succulent,
orange then red imitation fruit. Seeds including the
Podocarpus ledermannii Pilg., Bot. Jahrb. Syst. 54: epimatium ovoid, 1013 810mm, with a distal,
210. 1916. Types: Papua New Guinea: West Sepik, barely elevated crest.
Sepik River, [Lordberg], C. L. Ledermann 9943,
C. L. Ledermann 10064a (syntypes K, lectotype not Distribution
designated).
Papuasia: New Guinea, Bismarck Archipelago.
Podocarpus idenburgensis N. E. Gray, J. Arnold TDWG codes: 43 BIS NWG-IJ NWG-PN
Arbor. 39: 447. 1958.
Ecology a rounded crown in large trees. Foliage branchlets
slender, terete, finely grooved between remote leaf
Podocarpus ledermannii occurs as a scattered and bases. Terminal buds 39mm long, outer scales
locally common tree in tropical evergreen rainfor- lanceolate, terminating in an acuminate, slightly
ests, from near sea level to 2300m a.s.l. It has been spreading apex. Leaves on saplings and young
found in forests in river deltas (e.g. Purari, Sepik) as trees larger than on mature trees, 1222cm long,
well as in upland lower montane forests dominated ca. 15mm wide, acute, obtuse or truncated at apex.
by Castanopsis. It often grows in the understorey, but Leaves on mature trees linear-lanceolate, (5)611
can attain canopy height in medium-high forests or (14)cm long, (7)1014mm wide, straight or
in more open (disturbed?) forests. slightly falcate, narrowing at base to a 49mm long
880 petiole; margins often somewhat wavy; apex acute,
Conservation obtuse or narrowly rounded. Midrib on adaxial
(upper) side obtusely raised, especially near base,
IUCN: LC mostly less than 0.5mm wide and often fading
towards apex, raised and continuous on abaxial side,
Uses 0.51mm wide. Stomata small, in numerous inter-
mittent lines in two bands on either side of abaxial
This species is relatively uncommon and is prob- midrib. Pollen cones axillary, solitary or in clus-
ably not distinguished from the more common spe- ters of 23, sessile, with several basal, carinate bud
cies like P. neriifolius. Therefore it can be assumed scales, cylindrical, elongating to 5060(80?)mm,
to be logged when occurring in accessible forest 23.5 mm wide; microsporophylls triangular,
and attaining suitable size. The uses of the wood are spreading, with two globose pollen sacs. Seed cones
similar to that of P. neriifolius, i.e. primarily light axillary, solitary on a short peduncle; receptacles
construction and carpentry, and in the delta areas subtended by two 24mm long bracts (foliola),
where it occurs boats, oars, masts, and spars. As far consisting of an axis with 12 fertile bracts and 12
as known this species is not in cultivation. sterile ones, fusing and swelling, initially 56mm
long, becoming ca. 8 6mm and red when ripe.
Seeds single, sometimes two on a receptacle, includ-
Podocarpus levis de Laub., Blumea 24 (2): 496. ing the epimatium 811(13?)mm long, 78mm
1979. Type: Indonesia: Papua, Yapen Island, wide, ovoid-globose with a rounded distal end. Seed
Mariatu, L. J. van Dijk bb 30484 (holotype L). proper not observed.
The species epithet means with a smooth surface; it This species is rather similar to P. neriifolius, from
refers to the upper surface of the leaves from which which it is distinct in its often bluntly rounded leaf
the midrib scarcely protrudes. apices. That character is, however, variable even on
the same branch, with truncate and acute leaf api-
Vernacular names ces e.g. on C. B. Robinson 309 (K) from Amboina,
which was cited in the protologue of this species, but
marisa, sanru (Sulawesi); wasiwarare (Yapen Island, the variation was not commented upon. Podocarpus
New Guinea) levis does not have acuminate leaves, as seen in juve-
nile plants of P. neriifolius, and the midrib on the
Description adaxial side is only prominent near the leaf base and
soon flattens or even fades out towards the apex (at
Trees to 25(35?) tall, usually with a straight bole; least in sicco). Whether to recognize such leaf dif-
trunk d.b.h. 40cm or more. Bark smooth, thin, on ferences as character states of a distinct species is
large boles with narrow, longitudinal flakes or strips, a moot point; we need more and better material
light brown weathering grey; inner bark pinkish or to decide, and this species is here merely given the
reddish brown, fibrous. Branches spreading, forming benefit of doubt.
Distribution less ridged from decurrent leaf bases; terminating in
obtuse-conical buds with several elongated, spread-
Malesia: Borneo (Kalimantan Timur), Sulawesi, ing to recurved outer scales 410mm long. Leaves on
Maluku [Moluccas], Talaud Islands; Papuasia: New juvenile plants (including seedlings) similar to leaves
Guinea (Papua). on mature plants but sometimes longer, to 14cm.
TDWG codes: 42 BOR-KA MOL SUL 43 NWG-IJ Leaves on mature trees linear-lanceolate, 510(13)cm
long, 510(12)mm wide, straight or slightly fal-
Ecology cate, gradually widening from a petiolate base with
parallel sides for most of their length; apex acute or
Podocarpus levis is a scattered tree, locally common, sometimes obtuse. Midrib on adaxial (upper) side
in evergreen primary rainforest. It has been found to conspicuous but obtuse, sometimes obscure towards 881
grow from near sea level to 1650m a.s.l. It occurs on apex, on abaxial side flattened but well visible. Leaf
various substrates; in Borneo (one locality known) it colour lustrous green above, light dull green below.
was found on limestone. Stomata very small, forming numerous intermittent
lines in two bands on abaxial side. Pollen cones axil-
Conservation lary, nearly sessile or short pedunculate, solitary or
with 23 together, subtended by short, triangular bud
IUCN: LC scales, cylindrical, elongating unequally to 1525mm,
2.53.5mm wide; microsporophylls triangular, with
Uses an obtuse apex and bearing two partly hidden, sub-
globose pollen sacs. Seed cones axillary, solitary on
This species is a valuable timber tree and logged with slender peduncles 817mm long; receptacles sub-
other species where large trees occur and are acces- tended by two spreading bracts (foliola) 2.53.5mm
sible. The wood is used for general construction, long, consisting of an axis with 34 bracts, 2 often fer-
carpentry and furniture making. This species is not tile and spreading distally, swelling and amalgamat-
known to be in cultivation. ing to an 8 45mm large, succulent and red body.
Seeds 12, rarely 3 on distal part of receptacle, includ-
ing the covering epimatium 79mm long, obliquely
Podocarpus longifoliolatus Pilg., in Engler, ovoid, with a small distal crest fading at maturity.
Pflanzenr. IV.5 [18]: 79. 1903 [longefoliolatus]. Seed proper not observed.
Type: New Caledonia: Grande Terre, Province Sud,
Mt. Mou, J. A. I. Pancher s.n., 1870 (holotype P). Distribution
Podocarpus macrophyllus (Thunb.) Sweet var. pili- China: Anhui, Chongqing, Fujian, Guangdong,
ramulus Z. X. Chen & Z. Q. Li, Bull. Bot. Res. North- Guangxi, Guizhou, Hong-Kong, Hunan, Jiangxi,
East. Forest. Inst. 9 (3): 69. 1989. Yunnan, Zhejiang; N Myanmar [Burma]; S Japan;
Taiwan.
Description TDWG codes: 36 CHC-CQ CHC-GZ CHC-HU
CHC-YN CHS-AH CHS-FJ CHS-GD CHS-GX CHS-HK
Adult leaves (3)510(11) cm long, (5)710( CHS-HN CHS-JX CHS-ZJ 38 JAP-HN JAP-KY JAP-SH
13)mm wide, abruptly converging to an obtuse or TAI 41 MYA
apiculate apex, sometimes subacute.
Conservation
Distribution
IUCN: LC
China: Anhui, Chongqing, Fujian, Guangxi,
Guizhou, Hubei, Hunan, Jiangsu, Jiangxi, Sichuan,
Yunnan, Zhejiang; Japan: Honshu, Shikoku; Taiwan
(extreme S and Lanyu or Orchid Island).
TDWG codes: 36 CHC-CQ CHC-GZ CHC-HU
CHC-SC CHC-YN CHS-AH CHS-FJ CHS-GD CHS-GX
CHS-HN CHS-JX CHS-ZJ 38 JAP-HN JAP-SH TAI
Podocarpus madagascariensis Baker, J. Linn. Soc., glaucous at first, olive green, ripening to brownish
Bot. 21: 447. 1885. black. Seed proper not observed.
The species epithet refers to Madagascar, where this Baker (op. cit.) cited three collections as seen by him
species is endemic. in the protologue of this species: R. Baron 2794, R.
Baron 3129 and G. W. Parker s.n. All three are repre-
Vernacular names sented by specimens in the Kew Herbarium (K) and
are therefore syntypes. However, whereas the two
Hetatra, Tsindrodravina (Malagasy) collections made by Baron are sterile foliage rep- 887
resenting young trees (or a single tree), the Parker
Description specimen is an adult foliage branch and has a seed.
This latter specimen also bears a note from R. Baron
Trees to 25m tall, but often much smaller and (in litt. July 1884) about the uses of the wood and the
shrubby; trunk to 60cm d.b.h. Bark thin, exfoliating native name of the tree, which Baker also cites in the
in strips on larger trunks, in small flakes on small protologue. It is therefore appropriate to designate
trees, light brown weathering grey. Branches numer- the specimen at K collected by G. W. Parker as the
ous, spreading; foliage branchlets spreading or lectotype of Podocarpus madagascariensis Baker.
ascending, stout, terete, finely ridged and grooved,
terminating in robust, subglobose buds 46mm Distribution
wide, with imbricate, rounded to ovate scales; lower
scales carinate and weakly acuminate, with scari- Madagascar, along eastern plateaus and mountains.
ous upper margin. Leaves highly variable in size, on TDWG codes: 29 MDG
the type specimen 36cm long, 57mm wide, but
ranging from 218cm long and 316mm wide on Ecology
other specimens, elliptic-oblong to linear, mostly
thick coriaceous, rigid, but in one variety more lax Podocarpus madagascariensis occurs in moist forests
and drooping, tapering towards a petiolate base and or forest remnants from level plains near sea level
to an acute, acuminate (or long attenuate) or obtuse to forested ridges and escarpments at 20002400m
apex; margins slightly revolute; leaf colour lustrous a.s.l. In the lowland forest it can attain 2025m and
green above, dull green below. Midrib inconspicu- become a canopy tree with mostly evergreen tropical
ous or obtuse on adaxial (upper) side, continuous or angiosperms; on exposed rocky ridges it is a shrub or
petering out distally, more distinct and continuous, a stunted tree occurring in sclerophyllous low forest
with abruptly raised edges, on abaxial (lower) side. or scrubland rich in lichens, often on relatively dry
Stomata in numerous intermittent lines on either sites. The drooping leaved variety procerus occurs
side of midrib on abaxial side. Pollen cones axillary, from near sea level to about 1000m a.s.l. on sand or
solitary or with 23 together on short peduncles, sandstone; the small leaved var. rotundus occurs at
subtended by imbricate, rounded bud scales, cylin- altitudes between 1500m and 1800m a.s.l. on igne-
drical, elongating to 1.52.5(3)cm long, 45mm ous rock outcrops. The variety madagascariensis is
wide; microsporophylls imbricate, with triangular, much more widespread and occurs on diverse sub-
minutely denticulate apex, each bearing two basal, strates, usually above 800m a.s.l.
oblong pollen sacs. Seed cones axillary, solitary on
1523mm long, slender peduncles, consisting of an Uses
axis with 23 fused bracts which retain free bract tips
at distal widest end, only swelling slightly to a 45 mm According to one of the collectors of the original
long receptacle (sometimes virtually absent). Seeds material of this species, the Reverend R. Baron, its
large, ovoid-globose or nearly globose, including the wood was extensively used in house building, for
epimatium 1523mm long , smooth, weakly crested, flooring, etc. in the latter decades of the 19th century.
These uses have diminished with many of the larger Conservation
and more accessible trees now logged. The species
and its varieties are not recorded in cultivation. This variety was listed as Endangered in a pre-1994
IUCN Categories and Criteria listing on the grounds
3 varieties are recognized: of its rarity and restricted distribution near Tolanaro
[Fort Dauphin] in the SE of Madagascar. However,
it is also known from the Massif de Bekolosy in
Podocarpus madagascariensis Baker var. the far north of the island, and from a locality in
madagascariensis. Type: Madagascar: [central Fianarantsa Province. Logging and deforestation are
Madagascar], G. W. Parker s.n., Aug 1880 the main threats.
888 (lectotype K, here designated). IUCN: EN [B2ab (ii, iii, v)]
Description
Podocarpus madagascariensis Baker var. rotundus
Leaves coriaceous, more or less rigid and spread- L. Laurent, Ann. Fac. Sci. Marseille 23: 59. 1915.
ing, (2)315(18)cm long, (3)514(16)mm wide; Type: Madagascar: Massif du Manongarivo, 1500
apex acute or obtuse. Seeds including the epimatium 1800m alt., J. M. H. A. Perrier de la Bathie (P, not
ovoid-globose. designated).
Distribution Description
Madagascar, along the eastern plateaux and Leaves (3)57cm long, 46mm wide, acuminate
mountains. or with a long attenuate apex. Seeds including the
TDWG codes: 29 MDG epimatium nearly globular.
Ecology Description
Podocarpus micropedunculatus is known to occur Trees to 35m tall; trunk to 1.8(3.5) m d.b.h., but
as an understorey tree in Agathis forest, where it is small and stunted at highest altitudes. Bark thin,
accompanied by other podocarpaceous conifers and smooth or shallowly fissured longitudinally, exfoli-
by angiosperms adapted to poor sandy soil (keran- ating with small, thin flakes or strips, light or dark
gas forest). It is more abundant and may form a brown, weathering grey. Branches spreading, form-
major element in the vegetation in shrubby thickets ing a broadly domed crown in large trees. Foliage
along the margins of forest clearings, again mostly on branches terete, finely grooved or ridged, in slow
sandy, podzolic soils, or on sandstone. It also occurs growing (exposed) crowns with old leaf markings,
in peaty soils, usually podzolic, of raised beaches, terminating in 34mm wide, obtuse buds with a
and in peat-swamp forest, from sea level to around few spreading and recurving outer scales. Leaves
500m a.s.l. Sometimes this species is associated on seedlings and young trees similar to those on
with Dryobalanops rappa and Shorea albida (both mature trees, but variable in size mostly in rela-
Dipterocarpaceae), again as an understorey tree. This tion to exposure and/or altitude, 1.55cm long on
species shares the rare habit of rhizomatous propaga- small, stunted trees above 3000m altitude, 517cm
tion with Podocarpus drouynianus in SW Australia; long on trees in forests at lower altitudes, 516mm
the latter however never develops into a tree. wide, elliptic, lanceolate-linear to linear, straight or
slightly falcate, coriaceous; margins flat or slightly
Conservation revolute, gradually tapering down to a petiolate base
and to an acute apex. Midrib raised at least from
This species is insufficiently known and since its leaf base to beyond the middle, situated in a shallow
description and naming in 1985 few collections have groove, ca. 0.5mm wide on the adaxial (upper) side;
been made, so that an assessment using herbarium continuous to apex, wider and flattened on abaxial
based data was not possible. Its growth habit and side. Leaf colour lustrous green above, dull green
occurrence on the edges of forest clearings seem to or sometimes glaucous green below. Stomata small,
indicate that it may have declined but is probably not in numerous intermittent and wavy lines on either
in danger of extinction. side of abaxial midrib. Pollen cones axillary, solitary,
IUCN: NT (A2) sessile or nearly sessile, subtended by imbricate,
broadly rounded, carinate bud scales, cylindrical,
Uses elongating to 3040mm, 33.5mm wide at anthe-
sis; microsporophylls spirally arranged, imbricate,
No uses have been recorded for this shrubby species. with slightly elongated apex and minutely erose-
It could perhaps be suitable as a shrub in tropical to denticulate margin, bearing two basal pollen sacs.
subtropical gardens. Seed cones axillary, solitary on slender ca. 10mm
long peduncles, the two bracts (foliola) at base ravines. In eastern Africa common associated angio-
small and early deciduous; receptacles consisting of sperm tree genera in the forests are Albizia, Croton,
an axis with two bracts fusing and swelling to a 1013 Macaranga, Ocotea, Olea, Schefflera, and Syzygium;
79mm, succulent red body. Seeds normally soli- in Cameroon Agauria, Albizia, Ilex, Nuxia, Olea,
tary at oblique distal end of receptacle, including the Pyseum, Schefflera, and Syzygium are common with
epimatium ca. 8 10mm, ovoid-globose without a P. milanjianus, e.g. on Mt. Oku. Elfin forest occurs at
distal crest, green or whitish pruinose, turning dark the highest altitudes above ca. 2800m and can con-
purple. Seed proper ca. 6 8mm with a smooth, sist largely of Erica arborea in eastern Africa, or is
hard seed coat, brown. mixed, with e.g. Agauria, Cussonia, Myrica, Myrsine,
Philippia, Syzygium, and Tecomaria. Interfaces with
Taxonomic notes grassland (Arundinaria alpina) occur here, often 893
with poor drainage and P. milanjianus concentrated
According to the protologue, the type should be along rocky streams.
from original material cited as Habitat in Milanji,
6000 ft, Oct. [1891] Nos. 34 & 39 of Alexander Conservation
Whytes collections, which are kept at BM. In that
herbarium, the only original material now extant is IUCN: LC
a specimen collected by Whyte in October 1891 on
Mt. Mulanje, but without a number mentioned on Uses
the original sheet fragment; it was separated from
another persons collection and remounted, stating it This species is an important timber tree in many
is part of Type. It is here designated as the lectotype parts of tropical Africa. Its wood is valued for car-
of Podocarpus milanjianus. pentry and joinery as it is light coloured, even
grained, easily worked, and large trees yield good
Distribution sizes of sawn timber. More specialized uses requir-
ing high grade timber are veneer, furniture making,
Tropical Africa: Angola, Burundi, Cameroon, cabinet making, interior trim, household utensils,
Congo Republic, Kenya, Malawi, Mozambique, and wood carving. It has been used in afforesta-
Nigeria, Rwanda, S Sudan, Tanzania, Uganda, Dem. tion on a small scale in several African countries,
Rep. Congo [Zaire], Zambia, Zimbabwe. within and perhaps without its natural range. It is
TDWG codes: 22 NGA 23 BUR CMN RWA ZAI not known to be used in horticulture and is probably
24 SUD 25 KEN TAN UGA 26 ANG MLW MOZ ZAM restricted to a few botanic gardens in Africa and/or
ZIM other tropical countries or in glasshouses in cooler
regions. Provenances from high altitudes may well
Ecology be hardy enough to survive in European and other
countries with mild winters and abundant rainfall,
Podocarpus milanjianus is a montane to high mon- but no one seems to have tried it.
tane species occurring in tropical evergreen rain-
forest, cloud forest, or at its highest limit in dwarf
forest dominated by Ericaceae, interfacing with Podocarpus nakaii Hayata, Icon. Pl. Formos. 6:
subalpine grassland. The altitudinal range is (900 66. 1916. Podocarpus macrophyllus (Thunb.) Sweet
)13003000(3250) m a.s.l. and about as great in var. nakaii (Hayata) H. L. Li & Keng, Taiwania
west tropical as in east tropical Africa. It does not 5: 39. 1954. Type: Taiwan: Nantou Co., Holisha,
form conifer forest, although it can occur in forest Ta-shu-ku, [Toshoko], S. Nakai s.n. (holotype TI).
dominated by Juniperus procera in e.g. Kenya, and is Fig. 276, 277
usually accompanied by angiosperm trees; however
it will often become more abundant on exposed, Etymology
thin-soiled mountain ridges where the forest
becomes low and, at high altitudes, even stunted. In This species was named after the Japanese plant col-
tall forest it is often restricted to stream sides in deep lector S. Nakai.
Vernacular names forests at low to middle elevations along the steep
eastern coast of the island.
tai wan luo han song (Chinese)
Distribution
Description
Taiwan (Chianghua Co.?, Nantou Co., Taichung Co.).
Trees to 20m tall, with erect, terete trunk. Bark not TDWG codes: 38 TAI
described or observed. Foliage branchlets terete,
grooved, glabrous, terminating in globose buds ca. Ecology
4mm diam. with imbricate, triangular scales. Leaves
894 lanceolate to linear-lanceolate, (3)510(12) cm Podocarpus nakai occurs scattered in broad-leaved
long, 815mm wide, straight or slightly curved (angiosperm) forests in the central mountains of
towards apex, tapering gradually to a petiolate and Taiwan. These forests are dominated by the fami-
slightly twisted base ca. 5mm long and an obtuse or lies Fagaceae and Lauraceae, with Theaceae and
sometimes acute apex; margins entire and smooth; Magnoliaceae also prominent. Characteristic genera
midvein acutely raised adaxially, slightly raised or of trees are Cyclobalanopsis (= Quercus), Lithocarpus,
nearly flat abaxially; leaf colour mid green above Machilus (= Persea), Cryptocarya, Schima, and
(adaxially), pale green below; flushing leaves red- Magnolia. The altitudinal range of this species rich
dish pink or sometimes light green. Stomata on evergreen forest type is from 700 to 1800m a.s.l. and
abaxial side, very small, in numerous irregular lines it exists under a markedly monsoonal subtropical
on either side of midrib. Pollen cones axillary, ses- climate.
sile, solitary or in clusters of 23, becoming long
cylindrical and lax at anthesis, 2040mm long, Conservation
23mm wide; microsporophylls arranged in steep
spirals, triangular-apiculate, spreading, yellowish This species although not specifically targeted by
green, with two sublateral, globular, yellow pollen loggers as it is generally of only small to moderate
sacs. Seed cones axillary, solitary on 210mm long size, has suffered greatly from general deforestation
peduncles; receptacles subtended by (1)2 small, in the period after World War II. Much of the pri-
more or less linear bracts (foliola), 1014mm long, mary forest where P. nakaii occurred has gone, or it
swelling to 10mm thick, succulent, changing colour has been turned into secondary vegetation unsuit-
from glaucous green to yellow to crimson red when able for this species. A locality in Chianghua where
ripe, subtended by two small, leaf-like, deciduous W. R. Price collected the species in 1912 probably no
bracts. Seeds solitary, obliquely ovoid, 89 6mm, longer has it growing there; Flora of Taiwan, ed. 2,
slightly flattened, covered by a pruinose epimatium, 1: 565 (1994) only mentions Nantou for its distribu-
ripening blackish blue, crested distally. Seed proper tion. Fewer than 250mature trees are now thought
ovoid, 7 5mm. to exist.
IUCN: EN (A1a, D)
Taxonomic notes
Uses
Podocarpus nakaii is similar to P. macrophyllus and
has been treated as a variety of it. The pollen cones No uses have been recorded for this species; its
are solitary or with 23 together, not in clusters of wood was logged with other trees, and if of good
35 as in P. macrophyllus, and the leaves are usually size and shape would have been used for construc-
wider, but the ranges of these measurements overlap. tion, carpentry, etc. It is probably in cultivation, but
Podocarpus macrophyllus var. macrophyllus is culti- it may masquerade under the name Podocarpus
vated in Taiwan, but its var. maki occurs in natural macrophyllus.
Podocarpus neriifolius D. Don, in Lambert, Descr. with several basal, carinate bud scales, cylindri-
Pinus 2: [21]. 1824. cal, elongating to 2540(50)mm, 23.5mm wide;
microsporophylls spirally arranged, triangular,
Etymology spreading, with two globose pollen sacs. Seed cones
axillary, solitary on a 520mm long peduncle; recep-
The species epithet means leaf like that of the genus tacles with 2 subulate bracts (foliola) 26mm long
Nerium [oleander]. at base, swelling to become ellipsoid with a truncate
distal end, 810mm long, yellow turning orange-red
Vernacular names or bright red and succulent when ripe. Seeds solitary
at truncate end of receptacle, enclosed in a smooth
Brown pine; podo bukit (Malay); numerous local epimatium, green turning purplish green when 895
names are in use in Malesia (see De Laubenfels, ripe, ovoid-oblong or ovoid-globose, (8)1015
Flora Malesiana, ser. 1, 10 (3): 401. 1988); bai ri qing 78mm. Seed proper ovoid, 610mm long, slightly
(Chinese); Thng tre l di (Vietnamese). Pilger flattened.
(1903) cites kimerah and kiputri as names for the
Indonesian island of Jawa, but these seem to apply to Taxonomic notes
several species of podocarps now in different gen-
era. In Vanuatu the name nuhuoto was recorded; in In Flora of China 4: 83 (1999) and in the World
Fiji (Viti Levu) asimbolo, (Vanua Levu) kuasi. Checklist and Bibliography of Conifers (Farjon,
1998, [2001]) P. annamiensis, described by N. E.
Description Gray (op. cit.) from Indochina, Myanmar [Burma]
and Hainan Island, is recognized as a distinct spe-
Shrubs or trees to 45m tall, to 75(100)cm d.b.h., cies, in China restricted to Hainan. However, the
bole straight, in large trees slightly fluted or rarely stated differences with P. neriifolius are very small
buttressed at base. Bark smooth, thin, on large boles and seem to be inconsistent. It appears that these dif-
with narrow, longitudinal flakes or strips, light ferentiating character states (e.g. apiculate leaf apex,
brown weathering grey; inner bark pinkish or red- conspicuously raised midvein on both sides of the
dish brown, fibrous. Branches spreading, forming a leaf in P. annamiensis) can be found in P. neriifolius
rounded or domed crown. Foliage branchlets terete, from parts of its range beyond the areas where P.
more or less grooved, glabrous, terminating in 25( annamiensis is supposed to occur. Podocarpus anna-
8)mm long buds with spreading, triangular to lan- miensis is therefore here treated as synonymous, in
ceolate outer scales. Leaves on juvenile plants short agreement with recent treatments of the two taxa
petiolate, linear-lanceolate, 1520(25) cm long, especially concerning the conifers of Viet Nam
1726mm wide, often curved and with undulating (Nguyen Tien Hiep & Vidal, 1996; Nguyen Tien Hiep
margins, abrubtly narrowed at base, long acumi- et al., 2004). See also the Taxonomic notes under P.
nate or acute. Leaves of mature trees usually much subtropicalis for an opposite view on which taxa to
shorter and narrower, shade leaves (6)815cm exclude from or include with the widespread and
long, (8)1218mm wide, acuminate; leaves more variable species P. neriifolius. Gray (1958) described
exposed to light mostly linear-lanceolate or lanceo- P. neriifolius var. degeneri from Fiji (Viti Levu) as a
late, (4)812(15)cm long, 615mm wide, straight distinct variety. Silba (1984) elevated it to a species,
or slightly curved, petiolate at a more gradually while De Laubenfels (1985) separated the two species
narrowing base, tapering to an acute apex. Midrib into different sections, apparently based on slightly
acutely (sometimes obtusely) raised adaxially, flat or different sizes of the foliola or basal bracts on the
obtusely (sometimes acutely) raised abaxially, usu- receptacles. Such minutiae hardly justify classifica-
ally narrower on adaxial side; leaf colour dark green tions at the species level, considering variations, and
above, pale green below; new leaves flushing red. are not evident in the specimens seen at Kew (K).
Stomata very small, in numerous irregular lines on In a revision of Fijiian conifers, Doyle (1998) has
abaxial (under) side on either side of midrib. Pollen treated P. neriifolius var. degeneri as a synonym of
cones axillary, solitary or in clusters of 23, sessile P. neriifolius. The narrower leaves appear to justify
varietal distinction and I follow Netta Gray in this Myrtaceae, Sapindaceae, Guttiferae (Clusiaceae),
case. Yet another new species, P. epiphyticus, was Cunoniaceae, Rutaceae, and many others.
described from Myanmar [Burma] by De Laubenfels
and Silba (1988). In the protologue it is said to be Conservation
an epiphytic shrub which was taken from the type
specimen label. The isotype specimen (J. Keenan This species, while logged extensively as a valuable
3081 at K) is a branchlet with typically broad juvenile timber tree, is too widespread and common in most
foliage leaves belonging to P. neriifolius. It was prob- areas to be classified as threatened with extinction
ably a young plant germinated and growing upon a under the IUCN criteria. Locally, its status may be
tree, as conifers (and other trees) sometimes do, but more severe depending on rates of logging and lev-
896 it can be categorically stated that there are no obli- els of general deforestation. The species is listed on
gate epiphytes among conifers. CITES Appendix III (at the request of Nepal in 1975),
which means that certificates or permits are required
Distribution to export its timber from that country.
IUCN: status listed under varieties.
From E Nepal and Bangladesh through Myanmar
[Burma] and Indochina (including the Andaman Uses
Islands), S and SW China, Malesia, Papuasia,
Vanuatu and Fiji. This widespread species is an important timber
TDWG codes: 36 CHC-CQ CHC-GZ CHC-SC tree in several countries where it is common. Its
CHC-YN CHH CHS-FJ CHS-GD CHS-GX CHS-HN excellent wood is used for house construction and
CHS-JX CHS-ZJ CHT 40 ASS-ME ASS-MI BAN NEP carpentry and for making oars, spars, and masts
41 AND-AN CBD LAO MYA THA VIE 42 BOR-BR of sailing vessels. More specialized uses requiring
BOR-KA BOR-SB BOR-SR JAW LSI-BA LSI-ET high grade timber are veneer, furniture making,
LSI-LS MLY-PM MOL PHI SUL SUM 43 BIS NWG-IJ cabinet making, interior trim, household utensils,
NWG-PN SOL-NO SOL-SO 60 FIJ VAN and wood carving. In New Guinea, Highland tribes
use the bark to insulate walls of round houses. As
Ecology far as known it is not in cultivation; in China and
several countries where plants under this name
Podocarpus neriifolius is the most widespread species are introduced and/or cultivated, these com-
in the genus. It occurs as a scattered tree in evergreen monly belong to P. subtropicalis originating from
primary broad-leaved forests, either in the understo- Emei Shan [Mt. Omei] in Sichuan, China, or per-
rey or more commonly (and eventually) as a canopy haps to yet other similar species. This identity may
tree. Its altitudinal range in China is from 100m to also apply to a cultivar with variegated leaves that
1500m a.s.l., but in tropical Malesia it occurs from is occasionally seen labeled under P.neriifolius.
near sea level to 2900m a.s.l. It is usually confined
to well drained sites away from rivers, but occa- 2 varieties are recognized:
sionally it occurs in swampy forest. The soils vary
from nutrient-poor sands (kerangas in Borneo),
sandy clay or sandstone, to andesitic laterites (lato- Podocarpus neriifolius D. Don var. neriifolius.
sols) and ultrabasic soils over serpentine in New Margbensonia neriifolia (D. Don) A. V. Bobrov &
Guinea. In China and Indochina it is a component Melikyan, Bjull. Moskovsk. Obsc. Isp. Prir., Otd.
in Lithocarpus-Castanopsis mixed evergreen for- Biol. 103 (1): 60. 1998. Type: Nepal [Legi ad
est. In Malesia this conifer is commonly associated Sankoo,Napaliae 1821], N. T. Wallich 6052A
with other podocarps e.g. Dacrycarpus imbricatus, (lectotype BM, isolectotype K-W). Fig. 278, 279
Dacrydium spp. and Podocarpus spp., and occasion-
ally with Agathis spp. and/or Phyllocladus tricho-
manoides, especially on mountain ridges, and with Podocarpus neriifolius D. Don var. polyanthus
numerous angiosperm trees of the families Fagaceae, Wasscher, Blumea 4 (3): 455. 1941; Podocarpus polyan-
thus (Wasscher) Gaussen, Trav. Lab. Forest. Toulouse Podocarpus neriifolius D. Don var. degeneri N. E.
T. 2, 1 (2, 21): 191. 1976 (nom. inval., Art. 33.2). Gray, J. Arnold Arbor. 39: 467. 1958. Podocarpus dege-
Podocarpus decipiens N. E. Gray, J. Arnold Arbor. 36: neri (N. E. Gray) de Laub. ex Silba, Phytologia Mem.
204. 1955; Podocarpus neriifolius D. Don var. decipi- 7: 61. 1984; Margbensonia degeneri (N. E. Gray) A. V.
ens (N. E. Gray) Silba, J. Int. Conifer Preserv. Soc. 7 Bobrov & Melikyan, Bjull. Moskovsk. Obsc. Isp. Prir.,
(1): 35. 2000. Otd. Biol. 103 (1): 60. 1998. Type: Fiji: Viti Levu, Mba,
Podocarpus annamiensis N. E. Gray, J. Arnold Arbor. Nandarivatu, O. Degener DA 14272 (holotype A).
39: 451. 1958.
Podocarpus annamiensis N. E. Gray var. hainanensis Description
Gaussen, Trav. Lab. Forest. Toulouse T. 2, 1 (2, 21):
175, 210. 1976 (nom. inval., Art. 37.1). Shrubs or small trees. Terminal buds large, to 8mm 897
Podocarpus epiphyticus de Laub. & Silba, Phytologia long, ovoid, outer scales acuminate to long attenuate
64: 290. 1988. or apiculate. Leaves on mature trees 612cm long,
Podocarpus neriifolius D. Don var. penibukanensis 610mm wide, linear, acute. Leaves on juvenile
Silba, J. Int. Conifer Preserv. Soc. 7 (1): 34. 2000. plants to 18cm long and 17mm wide.
Podocarpus neriifolius D. Don var. staintonii Silba, J.
Int. Conifer Preserv. Soc. 7 (1): 36. 2000. Distribution
Podocarpus nivalis occurs at higher altitudes in the No common names have been recorded of this
sub-alpine to alpine scrubland and tussock grassland species.
of the New Zealand mountains. Its altitudinal range
varies from 800m to 2500m a.s.l. depending on local Description
climate and exposure. It is a species that commonly
creeps low over rocks, but when competing with tall Shrubs 12m tall, rarely a small tree to 6m tall;
grasses or in tall scrubland it may grow to a more densely branched, erect and rounded or more
e longated in small trees. Bark fibrous, light brown The leaf shapes of the two species are also said to
weathering grey. Foliage branchlets numerous, slen- differ, with more lanceolate leaves in P. beecherae
der, terete, glabrous with minute grooves between and more linear shapes in P. novae-caledoniae. They
leaf bases, terminating in small, ovoid or subglobose are claimed to occur in different habitats, with P.
buds with imbricate, appressed, triangular, weakly novae-caledoniae termed a semi-rheophyte while
keeled, 12mm long bud scales. Leaves on juvenile P. beecherae occurs in maquis on dry rocky sites.
plants longer than on mature plants, up to 12cm. Closer scrutiny of these stated differences in her-
Leaves on mature plants (2.5)48(10)cm long, barium specimens at K has shown that these distinc-
35(6)mm wide, linear, straight or slightly falcate, tions either do not hold or are at best unconfirmed
gradually or sometimes more abruptly widening (colour of receptacle). A collection not known to
from a petiolate base, mostly with two parallel sides De Laubenfels from the le des Pins (Pic NGa) was 899
up to a gradually tapering, acute or obtuse apex. found away from any stream in maquis and had red
[Shortest leaves may differ in being either narrowly receptacles. The yellowish green receptacles found at
lanceolate and acute or sometimes slightly wider the type locality of P. beecherae could well be a fully
above the middle and then obtuse.] Midrib on adax- grown but yet unripe stage [when I visited the site in
ial (upper) side often running in a shallow V-shaped November 2005 the small receptacles were glaucous
depression, forming a groove or a faint raised rib but immature]; this is seen in several other species
only proximally, on abaxial side obtusely raised; leaf in the genus, where the colour turns from yellowish
texture coriaceous; leaf colour lustrous light green green to yellow, orange, red and even purple even-
above, dull light green below. Stomata numerous, tually. Receptacle colour is not a sufficient charac-
small, in two wide bands of intermittent lines on ter to base a species on. Besides, a mature greenish
abaxial side. Pollen cones axillary, sessile, solitary or yellow receptacle would be unlikely in the genus; its
23 together, narrowly cylindrical, 718mm long, colour is an adaptation to bird dispersal and there-
22.5mm wide at anthesis, yellow turning brown; fore has to be quite distinct from the foliage when
microsporophylls very small, triangular, with two the seed is ripe to be effective. The leaf shapes of
sublateral, globose pollen sacs. Seed cones axil- the type of P. beecherae and other collections cited
lary on slender peduncles 515mm long, solitary; by De Laubenfels fall entirely within the varia-
receptacles subtended by two small, recurved bracts tion seen in those specimens of P. novae-caledoniae
(foliola) and consisting of an axis with 23 fertile he did not include in his new species. Podocarpus
bracts which amalgamate and become swollen and beecherae is therefore considered a synonym of P.
succulent, 912mm long, 68mm wide, yellowish novae-caledoniae.
green to orange or (dark) red. [In some populations
receptacles seem to stay greenish but it is not clear Distribution
if they do not become more coloured eventually.]
Seeds solitary or two on a receptacle, including the New Caledonia: Grande Terre (Province Sud), le
epimatium irregularly ovoid, 68 56mm, with des Pins
or without a small crest, whitish green or pruinose, TDWG codes: 60 NWC
ripening to light brown, smooth and lustrous. Seed
proper not observed. Ecology
Distribution Etymology
S Mexico; Central America; Bolivia; Colombia; The species epithet refers to the island of Palawan,
Equador; Peru; Venezuela. Philippines.
Vernacular names much of which has occurred in recent decades. It is
not known if this has affected the type location, and
No common name has been recorded for this species. thereby this species, but assuming that it was present
in places outside the type location, it is likely to have
Description suffered serious decline. This species is not within a
protected area.
Trees to 7m tall. Buds on vegetative branchlets IUCN: CR [B1ab (iii)]
46mm long and 45mm wide at their base, with
triangular, erect or slightly spreading outer scales. Uses
Leaves linear-lanceolate, 10.518.5cm long, 811mm
wide, narrowing to a 57mm long petiolate base No uses are specifically known of this species. 903
and an acute apex.. Midrib on adaxial (upper) side Presumably it will have been used or is being used
acutely raised. Pollen cones axillary, sessile, solitary, for firewood locally.
cylindrical, 3545mm long, 6.58mm wide; micro-
sporophylls with a 4mm long, lanceolate apex and
two basal pollen sacs. Seed cones not observed. Podocarpus pallidus N. E. Gray, Bishop Mus. Bull.
[This brief description has been translated from 220: 46. 1959. Type: Tonga: Tongatapu Group, Eua,
the Latin description given in the protologue; no Ohonua, [western side of plateau region], H. E.
other information being available.] Parks 16212 (lectotype F).
This species was first described and named in 1988, The species epithet means pale and probably refers
based on a single collection, C. E. Ridsdale SMHI to the pale green leaves.
1502 (holotype L, isotype K) with aberrant pollen
cones. This is unsatisfactory and a critical compari- Vernacular names
son with other species in the region is needed, but
hampered by the lack of collections particularly uhiuhi (Tongan name)
from Palawan. It is a species that has here been given
the benefit of the doubt, but may be synonymous Description
with P. polystachyus.
Shrubs or small trees to 10m tall, usually monopo-
Distribution dial or branching low; trunk to 25cm d.b.h. Bark
light brown weathering grey; inner bark reddish
Philippines, Palawan (Pagdanan Range). brown. Foliage branchlets ascending or spread-
TDWG codes: 42 PHI ing, terete, with fine grooves or ridges, terminat-
ing in small ovoid-conical buds with 26mm long,
Ecology spreading or recurving and acuminate outer scales.
Leaves on juvenile plants 1015cm long, 1217mm
Closed tropical evergreen rain forest dominated by wide. Leaves on mature plants linear-lanceolate
angiosperms. to linear, (2.5)412cm long (very variable on the
same branch), 511mm wide, straight or slightly
Conservation falcate, gradually tapering to a petiolate base and to
an acute or acuminate, sometimes mucronate apex;
Not known from any other than the type loca- margins occasionally slightly revolute; leaf colour
tion, which is called Ibangley Brookside Hill in the lustrous light green above, pale green below. Midrib
Pagdanan Range at only 40m altitude. A map in the on adaxial (upper) side narrow (0.30.4mm wide),
Philippines Red Data Book showing the extent of acute but sometimes fading towards leaf apex, on
natural forest cover on the island of Palawan at pres- abaxial side wider (to ca. 1mm) and obtuse or flat-
ent demonstrates an extreme rate of deforestation, tened. Stomata small, in numerous intermittent
lines on abaxial side on either side of distinct mid- Podocarpus parlatorei Pilg., in Engler, Pflanzenr.
rib. Pollen cones axillary, solitary, sessile, subtended IV.5 [18]: 86. 1903. Podocarpus angustifolius Parl.,
by triangular bud scales, long cylindrical, 2540mm in Candolle, Prodr. 16 (2): 512. 1868, non Griseb.
long, 34mm wide at anthesis, pale yellowish white; (1866. Type: Bolivia: [locality not stated], T. Bridges
microsporophylls triangular, acute, with scarious s.n. (holotype not located, isotype K).
margins and two sub-basal pollen sacs. Seed cones
axillary, solitary on 515mm long, stout peduncles; Etymology
receptacles subtended by two ca. 2mm long, curved
bracts (foliola), and formed of an axis with 23 This species has been named after the Italian bota-
fused bracts of unequal size, swelling to a succulent, nist Filippo Parlatore (18161877).
904 red to blackish red body 1013mm long, ca. 10mm
wide at distal end. Seeds single, including the epima- Vernacular names
tium ovoid to subglobose, 1013 810mm, with or
without an obscure distal ridge, blue to brown with a pino del cerro, pino blanco (Argentina, Bolivia)
glaucous bloom. Seed proper not observed.
Description
Distribution
Shrubs or stunted trees to 12m tall, occasionally to
SW Pacific: Tonga (Eua and Uta Vavau). 20m tall; trunks of trees to 1.5m d.b.h. Bark slightly
TDWG codes: 60 TON fissured on larger stems, brown weathering grey.
Branches often ascending from near the ground,
Ecology but in closed forest trees have a clear bole to some
height, forming a broad crown. Foliage branches
Podocarpus pallidus occurs in tropical low forest, sub-verticillate, ascending to erect, slender, terete,
dominated by angiosperms. This species is usually with fine grooves running down from slightly
scattered in forest patches along the plateau escarp- elevated leaf bases, yellowish green turning light
ment and in ravines or gullies of two islands in the brown. Terminal buds broadly ovoid to subglobose,
Tonga Archipelago situated some 325 km apart. It 46mm long and wide, with numerous apiculate
has been found on limestone cliffs on Uta Vavau scales with minutely denticulate margins and free
and occurs from around 50m to ca. 250m a.s.l. tips, giving buds a bristly appearance; axillary buds
smaller, globose. Leaves on juvenile plants or sap-
Conservation lings larger than on mature plants, to 12cm long and
5mm wide. Leaves on mature plants patent to more
This species appears to occur on two islands, Eua or less erect, (2.5)48(9)cm long, 23mm wide,
and Uta Vavau, of the Tonga Archipelago. No col- narrowly linear, straight or slightly falcate, with a
lections are known from Tongatapu, Tofua or Late, short petiolate base; margins slightly revolute, very
other islands in the archipelago with low forested gradually tapering to an acute-pungent apex, thin
mountains and plateaux. Because this species is a coriaceous, green above, pale green below; flushing
minor component in broad-leaved tropical forest, it leaves light green. Midrib on adaxial (upper) side of
is assumed there are fewer than 500mature trees on leaves a shallow groove often fading towards apex,
each island. We have no information about possible on abaxial side a narrow ridge, continuous or raised
decline. This species is not known from any pro- in lower part of leaves only. Stomata very small, in
tected area. numerous intermittent lines on abaxial side. Pollen
IUCN: VU (D1, 2) cones in clusters of (3)46 or more due to corym-
bose branching, on axillary, slender, angular stalks
Uses 515mm long; individual cones axillary to a 23mm
long, lanceolate, acute bract, sessile and subtended
No uses have been recorded of this species. by small, triangular, carinate-apiculate bud scales,
cylindrical, 510mm long, 1.52mm wide; micro- unlikely to involve timber in any quantity that could
sporophylls triangular, with minutely denticulate be of commercial importance. This species rarely
margins and two globose pollen sacs. Seed cones attains tree sizes over 15m tall, but in Argentina trees
axillary, appearing with new leaves, solitary on from 1520m have been recorded. It is a species with
38mm long peduncles; receptacles composed of high ability to recruit. For example, it is common in
an axis with 23 fused, swollen bracts, 45mm long secondary forest along roads, abandoned farms, and
and red when mature. Seeds globose, including the grazing areas where the use of fire was diminished or
epimatium 56mm long, 45mm wide, with or halted. Deforestation of a general kind, in which this
without an inconspicuous crest. species, where it occurs as a tree, will be logged with
the rest, could be of conservation concern, as could
Distribution increased domestic use of its timber when alterna- 905
tive sources become scarcer.
Argentina (Catamarca, Jujuy, Salta, Tucumn); IUCN: NT
Bolivia (Chuquisaca, Cochabamba, La Paz, Santa
Cruz, Tarija). Uses
TDWG codes: 83 BOL 85 AGW-CA AGW-JU AGW-SA
AGW-TU The wood of this species is at present used locally
for firewood, fence posts and domestic articles, e.g.
Ecology simple furniture and tool handles. In Argentina it
was heavily used for paper pulp in the 1970s and
Podocarpus parlatorei occurs in montane scrubland 80s; now (1990s2000s) it is occasionally logged
and forest at altitudes between 1000m a.s.l. and the for local (domestic) uses only. In Bolivia there is a
upper limit of trees between 2400m and 3150m programme to exploit the species to use the wood
a.s.l. This species may grow into a tree of maximal for local housing.
20m height in riverine forest at lower altitudes, but
it is more often a shrub or stunted tree, branching
near the base. It is commonly associated in the for- Podocarpus pendulifolius J. T. Buchholz & N.
est with Meliaceae, Myrtaceae (typically with Alnus E. Gray, J. Arnold Arbor. 29: 138. 1948. Type:
-Betulaceae- in NW Argentina and S Bolivia) and Venezuela: Cordillera do Mrida, Paramo de la
other angiosperm trees; on cliff sides and in steep Negra, above La Grita, J. A. Steyermark 57115
rocky gullies it is often the dominant shrub and can (holotype F). Pl. 38
be quite abundant. In valleys it is locally abundant
on old river terraces or on river banks, often now in Etymology
forest relicts. It is a pioneer species with a key role
in the forest dynamics. Its seeds or fruits are a food The species epithet refers to the drooping or pendu-
resource for endangered and of restricted range bird lous leaves.
species, such as Penelope dabbenei (Cracidae) and
Amazona tucumana (Psittacidae). Vernacular names
Plate 38. Podocarpus pendulifolius; Podocarpus salicifolius. 1. Branch with leaves and seed cones. 2. Seed
cone. 3. Habit of tree. 4. Branch with leaves and pollen cones. 5. Seed cone. 12 = P. pendulifolius; 35 =
P. salicifolius.
turning light brown. Terminal buds subglobose, (sub)populations ca. 100 km apart. Trees of some
35mm wide, often shorter than wide, with imbri- size are known to be valued and used for building
cate, broadly ovate, carinate scales with scarious houses, so there must be logging targeted at this
margins. Leaves strongly drooping or pendulous; species. No (sub)populations have been reported in
leaves on the upperside of the shoot curved down protected areas.
above a petiolate base, 512cm long, lanceolate-lin- IUCN: EN [B1ab (ii, iii, v), B2 ab (ii, iii, v)]
ear, 711mm wide, straight or slightly falcate, gradu-
ally tapering at both ends; margins nearly flat; apex Uses
acute; texture thin coriaceous; leaf colour green on
both sides. Midrib obtusely raised and continuous The wood of this species is considered to be of good
to apex on both sides of leaf. Stomata very small, quality for construction of houses. The pendulous or 907
in wavy, intermittent lines on abaxial side in broad drooping habit of the foliage would make an attrac-
bands on either side of midrib. Pollen cones axil- tive feature for horticulture and provenances from
lary, solitary, sessile, with obtuse perular scales at the highest altitudes may even prove more or less
base, cylindrical, 2040mm long, 67mm wide at hardy. It is not known to be in cultivation.
anthesis; microsporophylls broadly ovate, obtuse
with erose-denticulate margin, bearing two globose
pollen sacs. Seed cones axillary, solitary on 710mm Podocarpus perrieri Gaussen & Woltz, Bull. Soc.
long peduncles; receptacles 89mm long, swelling Hist. Nat. Toulouse 111: 319. 1975. Podocarpus
and becoming red to dark purple, often pruinose. rostratus L. Laurent var. perrieri (Gaussen & Woltz)
Seeds single, attached at distal end of receptacle, Silba, J. Int. Conifer Preserv. Soc. 7 (1): 39. 2000.
including the epimatium 89mm long, 67mm Type: Madagascar: Antananarivo Prov., Toamasina,
wide, ovoid-globose, glaucous-pruinose, with a Fort dAndasib, J. M. H. A. Perrier de la Bathie
small apiculate crest or nearly obtuse. Seed proper 17109 (holotype P).
not observed.
Etymology
Distribution
This species was named after Henri Perrier de la
NW Venezuela: Cordillera de Merida. Bathie, who collected the type specimen.
TDWG codes: 82 VEN
Vernacular names
Ecology
No common names have been recorded for this
Podocarpus pendulifolius occurs in montane mixed species.
low-canopy rainforest at elevations between 1400m
and 3000m a.s.l. At the highest elevations it is part Description
of a dwarfed forest on slopes immediately below
the paramo vegetation, an alpine grassland rich in Trees to 30m tall; trunk to 80cm d.b.h. Bark thin,
herbs. Young trees are said to be locally abundant becoming scaly on large trunks, light brown, weath-
in this habitat. It is less common in the taller forests ering grey. Branches ascending and spreading, in old
below these stunted forests, where it can occasion- trees forming an umbrella-shaped crown. Foliage
ally attain a height of 20m. branchlets numerous, ascending, slender, terete,
finely grooved between leaf bases, terminating in
Conservation small buds with imbricate, obtuse outer scales and
acute or apiculate inner scales with free but incurved
Based on distribution data from herbarium speci- apices. Leaves on saplings and young trees similar
mens, the extent of occurrence (EOO) is less than to those on mature trees, more remotely placed on
3000 km2 and the area of occupancy (AOO) is less branchlets and at the larger end of the size range.
than 100 km2; the species is only known from two Leaves on mature trees crowded, spreading to nearly
erect, 0.72(2.5)cm long, rigid, linear, straight or tion of tree size was made with the collection made
slightly curved, 12mm wide, irregularly oval in at 2000m a.s.l.
cross-section, narrowing slightly to a decurrent
base; apex pungent. Midrib absent or a very shal- Conservation
low groove on adaxial (upper) side, an obtuse keel
on abaxial side. Stomata small, in intermittent white This species is only certainly known from the type
lines forming narrow bands on either side of abaxial location and one other location and from GIS data
midrib. Pollen cones axillary, solitary or with 23 on it appears to be a deforested area. The type collec-
slender, 34mm long peduncles, with a few triangu- tion was made in 1925, the most recent collection
lar perular scales at their base, cylindrical, 815mm dates from 1951. Surveys in this area are urgently
908 long, 3mm wide at anthesis; microsporophylls with required to determine if this species is still extant.
an elongated, rostrate apex, bearing two ovoid- Deforestation is serious in this region and this spe-
globose pollen sacs at base. Seed cones axillary, cies is not known from any protected area.
solitary on short peduncles; receptacles subtended IUCN: CR [B1ab (ii, iii, v), B2ab (ii, iii, v)]
by two minute, deciduous bracts (foliola), consist-
ing of a short axis with two unequal bracts which Uses
become fused and coriaceous, 23mm long. Seeds
solitary with fertile bract, including the epimatium No commercial uses are recorded of this species, but
ca. 8 5mm, obliquely ovoid with a minute crest. it is undoubtedly used locally for firewood and, if
trees are large enough, for construction timber, such
Taxonomic notes as flooring of houses.
The species epithet means with many spikes and Indochina: southernmost peninsular Thailand;
refers to the clustered pollen cones. Malesia: Borneo, Peninsular Malaysia, Maluku
[Moluccas] (Obi & Waigeo Islands), Philippines,
Vernacular names Singapore, Sulawesi, Sumatera (Bangka & Belitung
Islands); Papuasia: New Guinea (Papua: Birds Head
podo laut (Malay); jati bukit (Peninsular Malaysia); Peninsula).
landin (Sarawak); kandabang (Sabah) and several TDWG codes: 41 THA 42 BOR-BR BOR-KA BOR-SB
more locally used names as listed in Flora Malesiana BOR-SR MLY-PM MLY-SI MOL PHI SUL SUM 43
ser. 1, 10 (3): 418 (1988). NWG-IJ 911
Description Ecology
Trees to 20(40) m tall, d.b.h. to 45cm (commonly This species occurs primarily on sandy beaches and
a monopodial or multi-stemmed small tree ca. bluffs, often on the land side of mangrove thickets at
6m tall). Bark shallowly fissured, peeling in long, or just above the high tide mark. It can even occur
fibrous strips, brown weathering grey; inner bark within the mangroves on slightly raised, sandy ridges
soft fibrous, pink. Branches spreading, forming an (probably old beaches). Here it is a stunted tree not
irregular to rounded and dense crown in most trees. exceeding 67m in height. On coastal limestone
Foliage branchlets terete, slightly grooved or ridged, and granitic rocks the trees are gnarled and shrubby,
terminating in 1.53mm long buds with narrowly not exceeding 23m. A second important habitat
triangular to lanceolate, erect to spreading outer is coastal lowland kerangas, a low forest on almost
scales. Leaves on juvenile plants linear to linear-lan- pure sand, and pandangs, i.e. degraded heath for-
ceolate, 711cm long, 813mm wide, more or less est; in these vegetations Myrtaceae have an impor-
abruptly tapering to a petiolate base and more grad- tant presence besides the conifers. In the interior of
ually to an acute or apiculate apex. Leaves on mature Peninsular Malaysia, the Philippines, and elsewhere
trees linear to linear-lanceolate or nearly oval when it occurs on limestone hills and plateaus at altitudes
very short, (2.5)511cm long, 611mm wide, short between 150m and 550m (in Palawan at ca. 1000m)
petiolate, abruptly narrowed at base, more gradu- a.s.l. Here trees may be taller; in Obi (Moluccas)
ally tapering to an acute or obtuse apex, straight or trees are reported to reach 40m tall with slender,
slightly curved, coriaceous; margins flat or nearly so; buttressed boles clear of branches to 25m. This can
midrib acutely raised and narrow adaxially (above), only occur in competition with other tall forest trees,
only slightly raised and wider abaxially (below); leaf where P. polystachyus is striving to reach the canopy.
colour deep green above, pale green below. Stomata It appears therefore to be a highly adaptable species
on abaxial side, very small and in numerous irreg- and the nearest to a mangrove-inhabiting conifer in
ular lines in two broad bands divided by midrib. existence.
Pollen cones clustered axillary with 25, sessile or
very short pedunculate, with a few small scales at Conservation
base, cylindrical to 1530mm long and 3mm wide;
microsporophylls imbricate, slightly spreading at This species although widespread is declining due to
anthesis, apiculate, with two oblong pollen sacs. Seed habitat loss both past and future in various parts of
cones axillary, solitary on a 26mm long peduncle; its range.
receptacles with 2 very small basal bracts to 1.5mm IUCN: VU (A4)
long (deciduous), 710mm long, swelling to 8mm
thick, pruinose, becoming orange-red to red when Uses
ripe. Seeds including the epimatium 79 57mm,
globose-ovoid, smooth, glaucous green. Seed proper Where this species grows into a tall tree, e.g. in
not observed. Maluku [Moluccas], it is a fairly important source
of podocarp timber. It has pale yellowish brown, gradually tapering to an acute apex; margins flat;
leight-weight wood used for construction, window midrib acutely raised and less than 0.5mm wide on
frames, boat building including masts, spars and adaxial side, obtuse and ca. 1mm wide on abaxial
oars, flooring, veneer, furniture making and cabi- (lower) side. Stomata very small and inconspicuous
net work, carpentry and joinery, household utensils, in dense lines on abaxial side. Pollen cones axillary,
matches, and toothpicks. Podocarpus polystachyus is solitary or occasionally in pairs, short peduncu-
one of few species in the genus commonly planted late, subtended by short obtuse bracts (bud scales),
as an ornamental tree in tropical countries, mostly elongating at anthesis to 3045mm, 34mm wide;
within its area of natural distribution but at least as microsporophylls with acute apex ca. 1mm long,
far away as the Congo Republic. bearing two globose pollen sacs. Seed cones axillary
912 on a 24mm long peduncle; receptacles subtended
by a 3mm long, thick bract (foliolum), 811mm
Podocarpus pseudobracteatus de Laub., Blumea 26 long, formed of an axis with two unequal bracts,
(1): 142. 1980. Type: Papua New Guinea: Southern swelling to become succulent and first orange, then
Highlands, Tari Subdistrict, Mt. Ambua, C. red. Seeds including the epimatium ovoid-globose,
Kalkman 5189 (holotype L). 1011 89mm when mature, distally obtuse. Seed
proper not observed.
Podocarpus archboldii N. E. Gray var. crassiramosus
N. E. Gray, J. Arnold Arbor. 39: 453. 1958. Distribution
Etymology Distribution
No vernacular names are recorded for this species, Podocarpus ridleyi occurs localized on several
which was originally described as a mere variety of mountain summits and ridges in somewhat stunted
the widespread P. neriifolius. rain forest; it may attain greater size when it occurs
in forest on slopes below these summits. Its altitu-
Description dinal range is from 480m to 1500m (possibly to
2100m) a.s.l. It grows in impoverished soils derived
Trees to 25m tall, in stunted forests less than 10m from sandstone or granite (Mt. Ophir) and can be
tall, d.b.h. to 40cm. Bark soft, flaky or scaly, pale the dominant tree in these localities.
Conservation ceolate (widest beyond the middle) to elliptic, on
juvenile plants somewhat linear and to 6cm long,
Rapid change in land use involving the conversion on mature plants 23(3.8)cm long, 47mm wide,
of rainforest into rubber tree plantations have had gradually narrowing to a short petiolate base, more
an impact on the trees that occurred on the lower abruptly ending in an acute or obtuse apex. Margins
slopes of at least the southern subpopulations, revolute; leaf texture coriaceous; leaf colour deep
reducing them to the higher summit areas. green above, pale or glaucous green below. Midrib
IUCN: VU [B2ab (ii, iii, v)] on adaxial (upper) side obtusely raised or nearly
flat with a central groove, on abaxial side promi-
Uses nently raised. Stomata on abaxial side very small,
916 in numerous intermittent lines on either side of the
No uses have been recorded of this species. midrib. Pollen cones axillary, solitary, sessile with
perular scales at base, cylindrical, 1520mm long,
2.53mm wide; microsporophylls triangular, with
Podocarpus roraimae Pilg., Notizbl. Bot. Gart. two globose pollen sacs. Seed cones axillary, solitary
Berlin-Dahlem 5: 299. 1913. Type: Guyana: Cuyuni on a 510mm long, slender peduncle; receptacles
Mazaruni, Pakaraima Mountains, Mt. Roraima, small, 68mm long, formed of a short axis with
E. H. G. Ule s.n. (holotype B). two unequal bracts, slightly swelling and ripening
red. Seeds single, including the epimatium ca. 8mm
Podocarpus buchholzii de Laub., Fl. Venezuela 11 (2): long, ca. 5mm wide, ovoid with a distal crest. Seed
31. 1982. proper not observed.
Podocarpus buchholzii de Laub. var. neblinensis Silba,
Phytologia 68: 66. 1990. Taxonomic notes
Podocarpus roraimae is found on isolated table Small, usually stunted trees 810m tall. Bark exfo-
mountains (tepuis), mostly on sandstone, at alti- liating in small strips and flakes, light brown weath-
tudes between 1800m and 2700m a.s.l. This spe- ering grey. Foliage branchlets slender, terete, finely
cies occurs in small stands of shrubs and trees in a grooved on vigorous shoots of young plants, ridged
mozaic vegetation of ombrotrophic bogs dominated between decurrent leaf bases on older, slow grow-
by grasses and herbs and interspersed low scrub and ing trees, terminating in small ovoid-globose buds
woodland. It also occurs in ravines near streams and with imbricate, apiculate scales. Leaves on young
on wet plateaux and summits with boggy vegetation plants linear, 2.56cm long, 23mm wide, straight
characterized by various ferns, pitcher plants and or curved downward. Leaves on mature trees nar- 917
orchids. Shrubs of this species occur in more open rowly lanceolate to linear, (1)1.53(3.6)cm long,
terrain, while in forest it can attain small tree size. It straight or sometimes slightly falcate towards apex,
can be associated with P. steyermarkii and P. tepuien- 12mm wide, only slightly narrowing to a sessile,
sis, which are distinct in having larger, respectively decurrent base; margins entire, mostly parallel but
smaller, leaves with a midvein groove on the adaxial in the distal part tapering to an acute apex. Midrib
side. There is high rainfall and frequent mist, and absent or inconspicuous on adaxial (upper) side
temperatures can fluctuate strongly from day to by a shallow central groove, continuous but flat or
night and depending on weather conditions. obtusely raised on abaxial side. Stomata on abaxial
side in several intermittent lines on either side of
Conservation midrib. Pollen cones axillary, solitary or in pairs,
rarely 3 on 16mm long stalks, subtended by a
This species, originally described from Mt. Roraima, few keeled bud scales, cylindrical, 1015(20)mm
has since been found in several isolated tepuis in long, 2.53mm wide; microsporophylls more or
Bolivar and Amazonas, Venezuela. Due to the remote- less triangular, with an elongated, 1mm long ros-
ness of this region and its mostly undisturbed mon- trate apex and two relatively large basal pollen sacs.
tane rainforests, it is believed that this species may Seed cones axillary, solitary, pedunculate, formed
occur on yet other mountains and is not threatened. of an axis with two unequal bracts, subtended
One known location is within Roraima National Park. by two spreading or recurved 1mm long bracts
IUCN: LC (foliola), fusing and slightly enlarging to a cune-
ate, 34 24mm, coriaceous receptacle with
Uses divergent bract tips. Seeds solitary, including the
epimatium ca. 14 8mm, obliquely ovoid, with a
No uses have been recorded of this species. small crest slightly below the apparent apex. Seed
proper not observed.
The species epithet means red and refers to the The species P. indonesiensis de Laub. & Silba
colour of the flushing leaves. (op.cit.), based on a few collections by P. J. Eyma in
Sulawesi and an additional specimen from Amboina Podocarpus rumphii Blume, Rumphia 3: 214. 1847.
(Maluku), is perhaps only a high altitude form of P. Margbensonia rumphii (Blume) A. V. Bobrov &
rubens in Sulawesi (25003000m a.s.l.) with shorter Melikyan, Bjull. Moskovsk. Obsc. Isp. Prir., Otd.
and thereby more elliptical leaves. In New Guinea P. Biol. 103 (1): 60. 1998. Type: Indonesia: Papua
rubens has been collected from as high as 2750m, a [Nov. Guinea], leg. ign. s.n. (holotype L, barcode
similar altitude as in Sulawesi. Shorter leaves tend L00050854). Fig. 287
to be elliptical, longer leaves linear-lanceolate, and
both occur on the same specimens (J. F. Veldkamp Podocarpus koordersii Pilg. ex Koord. & Valeton,
8318, K, L). The specimen from Amboina was said to Meded. Lands Plantentuin 68: 268. 1904;
be from around 1000m a.s.l. All other characters are Margbensonia koordersii (Pilg. ex Koord. & Valeton)
identical with P. rubens and P. indonesiensis is here A. V. Bobrov & Melikyan, Bjull. Moskovsk. Obsc. 919
considered a taxonomic synonym. Isp. Prir., Otd. Biol. 103 (1): 60. 1998.
Podocarpus philippinensis Foxw., Philipp. J. Sci.
Distribution 6: 149. 1911; Margbensonia philippinense (Foxw.)
A. V. Bobrov & Melikyan, Bjull. Moskovsk. Obsc.
Malesia: Borneo, Flores, Maluku [Moluccas] Isp. Prir., Otd. Biol. 103 (1): 60. 1998.
(Seram), Sulawesi, Sumatera, Timor; Papuasia: New Podocarpus rumphii Blume var. arbainii Silba, J. Int.
Britain, New Guinea. Conifer Preserv. Soc. 7 (1): 38. 2000.
TDWG codes: 42 BOR-KA BOR-SB BOR-SR LSI-LS Podocarpus rumphii Blume var. aruensis Silba, J. Int.
MOL SUL SUM 43 BIS NWG-IJ NWG-PN Conifer Preserv. Soc. 7 (1): 38. 2000.
Ecology Etymology
Podocarpus rubens is a constituent of lower montane This species was named in honour of Georg Eberhard
to high montane primary rain forests. Its lower alti- Rumphius (16281702), author of an early Flora of
tude is around 800m a.s.l. (but sometimes lower to Amboina.
150m) and it reaches optimum growing conditions
between that altitude and 1500m. At higher altitudes Vernacular names
it reappears on exposed ridges between 2000m and
3200m a.s.l. as a small, often stunted tree in mossy Numerous local names are in use for this species;
forest, especially in New Guinea accompanied by several are listed by region in Flora Malesiana ser.
Nothofagus spp. and other fagaceous trees. It is much 1, 10 (3): 416 (1988); in Maluku (Moluccas) and in
less common in lowland swamp forest, where it may Sabah it is called kayu china; in Vanuatu it is called
grow as an occasional tree with Dacrydium spp. nimsal (Sie language).
Conservation Description
Ecology
Podocarpus rusbyi J. T. Buchholz & N. E. Gray,
Podocarpus rumphii is a constituent of lowland to J. Arnold Arbor. 29: 134. 1948. Type: Bolivia: La
lower montane tropical rainforests, where it can be Paz, H. H. Rusby 2463 (holotype NY).
locally common. It is likely to be confused with the
similarly widespread but more ubiquitous species Etymology
P. neriifolius, which can have similarly large leaves
(but usually narrower) and differs in characters not This species was named after the botanist and plant
always observable in specimens. The altitude ranges collector Henry Hurd Rusby (18551940).
Vernacular names line. In Peru it was found near Machu Pichu in high
montane forest at 2900m a.s.l. In Bolivia, where
pino blanco (Bolivia) the species is much more common and widespread,
its altitudinal range is between 1500m and 3350m
Description a.s.l. It is commonly associated in the forest with
Meliaceae, Myrtaceae and other angiosperm trees; at
Trees to 25m tall; trunk to 60cm d.b.h. Bark on larger high altitudes it forms thickets with other dwarfed
trees becoming fissured and scaly, brown weathering trees and shrubs. It occupies similar habitats as
grey. Branches spreading or ascending, forming a P.parlatorei within the same altitudinal range, but in
broad open crown in the forest, more bushy in open Bolivia the two species are geographically separated.
terrain at high altitudes. Foliage branches slender 921
or stout, terete, grooved between remote leaf bases, Conservation
terminating in broad conical buds, on stout branch-
lets 57mm wide at base, with lanceolate-acuminate There is evidence of forest decline in the region, but
outer scales 510(16)mm long and spreading out, we cannot quantify its impact upon this species. This
the inner scales smaller and erect or converging, buds species appears to have a limited distribution within
on slender lateral branchlets smaller, with shorter Bolivia, with an estimated extent of occurrence
scales. Leaves on saplings and shaded branches (in (EOO) of 22,500 km2 and an area of occupancy
forest) larger than on mature trees and exposed small (AOO) below 2000 km and with one location in
trees, 49cm long, 610mm wide. Leaves on mature Peru at ca. 300 km distance. Two localities from
and exposed trees (2)35cm long, (4)58(10)mm where collections were made are within protected
wide, patent, elliptic to linear-lanceolate, widest in areas: Parque Nacional de Choquecamiri in Bolivia
middle part, gradually or sometimes more abrubtly and Machu Pichu National Park in Peru.
narrowing to a short petiolate base; margins slightly IUCN: VU [B2ab ii, iii, v)]
revolute, gradually tapering to an acute apex, coria-
ceous, deep green above, pale green below. Midrib a Uses
continuous narrow groove on adaxial (upper) side,
raised but narrow (0.4mm wide) and continuous No commercial uses are recorded of this species. Its
to apex on abaxial side. Stomata small, in numerous wood is used locally for construction purposes. This
intermittent lines on abaxial side. Pollen cones not species is not known to be in cultivation.
observed, presumably solitary. Seed cones axillary,
solitary on 310mm long peduncles; receptacles com-
posed of an axis with two unequal, spreading bracts, Podocarpus salicifolius Klotzsch & H. Karst. ex
fusing and swelling to 68mm long with exserted, Endl., Syn. Conif.: 209. 1847. Type: Venezuela:
triangular bract tips, green becoming red. Seeds sin- Cordillera de la Costa, Distrito Federal, Gran
gle between bract tips, ovoid-globose, including the Colombia, [Habitat in Colombia], H. Karsten s.n.
epimatium ca. 7mm long with an obtuse distal crest, (holotype W, destroyed, isotype LE). Pl. 38
dark green. Seed proper not observed.
Podocarpus pittieri J. T. Buchholz & N. E. Gray,
Distribution J. Arnold Arbor. 29: 130. 1948.
Distribution Description
Bolivia; NW Brazil; Colombia; Peru; N Venezuela. Trees to 20m tall; trunk monopodial or branching
TDWG codes: 82 VEN 83 BOL CLM PER 84 BZN low, up to 1m d.b.h. Bark thin, smooth in young
trees, becoming ragged with exfoliating strips on
Ecology older stems, light brown weathering whitish grey.
Branches in young trees forming a pyramidal crown,
Podocarpus salicifolius occurs in the Andes ranges in larger trees spreading out more widely, form-
in montane to upper montane rain forest or cloud ing a broadly domed, often dense crown. Foliage
branchlets slender, often drooping, terete, with fine plantation forest with Eucalyptus and Pinus radiata.
grooves between decurrent leaf bases, terminating in Increased fires have destoyed many other stands
small, globose buds with imbricate, ovate-triangular there. Cutting for firewood has also reduced the
scales. Leaves on seedlings and saplings similar to number of mature trees. In the south of its range it
those on mature trees, narrowly linear-lanceolate, is doing better, but there is a continuing decline of
straight or (slightly) falcate, (3)59(12)cm long, its habitat resulting in reduced extent of occurrence
(3)47mm wide, lax, narrowing to a short peti- (EOO) and area of occupancy (AOO) and, especially
olate base, gradually tapering to an acute apex, lus- in the north, fragmented subpopulations. Habitat
trous light green above, pale green below. Midrib on degradation is probably the most serious and wide-
adaxial (upper) face less than 0.4mm wide, raised spread menace to the survival of this species, which
from base but fading towards apex, similarly nar- does not react well to the opening up of forest cover 923
row but continuous to apex on abaxial side. Stomata and concomitant dehydration in summer.
very small, in numerous intermittent lines on abaxial IUCN: VU (A2a, c, d)
side. Pollen cones axillary, sessile, solitary or grouped
with two or more, often from flushing leaves, or from Uses
older leaves, 2550mm long, very slender, the longest
almost filiform, 1.52mm wide at anthesis, yellow; The fine-grained wood of this species is used locally
microsporophylls minute, triangular, bearing two in the craft industry and was in the past of impor-
globose pollen sacs. Seed cones axillary, solitary on tance for house building. More important now per-
slender, 12cm long peduncles; receptacles formed haps is its use in horticulture. Its date of introduction
of an axis with 23 fused bracts, one or two of these in Europe is uncertain, but it has been very success-
fertile, swelling to 56 45mm, succulent deep ful and is quite common in large gardens and parks
red to violet. Seeds solitary or sometimes two on a in regions with mild winters and abundant rainfall.
receptacle, including the epimatium 78 45mm, It usually forms a bushy tree with dense, pendulous
obliquely ovoid, green maturing to reddish, with an foliage.
obtuse crest. Seed proper not observed.
Ecology Etymology
In the north of its range (Mediterranean climate) The species epithet refers to the Solomon Islands,
Podocarpus salignus follows water courses in the where this species is endemic.
Roble-Hualo forest type (Bosque Maulino Costero);
in the Andes higher than 1000m a.s.l. it can form Vernacular names
pure stands in wetter areas away from water courses,
or it occurs in moist ravines. Here it is a minor com- dengali tolo (Malaita)
ponent of Nothofagus obliqua forests. The altitudinal
range of this species is between 20m and 1200m Description
a.s.l., with the highest localities in the Andes on
S-facing slopes. Trees to 25m tall, with a straight bole in forests to
35cm d.b.h., stunted on poor growing sites; trunk at
Conservation base not buttressed. Bark usually smooth, in larger
trees flaking with thin, curling strips, pale yellow-
In the northern part of its range there has been sub- ish brown; inner bark weakly fibrous. Branches
stantial decline due to conversion of natural forest to spreading in tiers, sinuous, often drooping, forming
a diffuse, conical crown, in old trees more irregu- low-lying small island, where it may have disap-
lar and rounded. Foliage branches terete, stout, peared. The other subpopulations are represented
grooved between leaf bases, terminating in a large by several herbarium collections from San Cristobal
conical bud with subulate-acuminate, keeled scales and one from Choiseul. Deforestation has mainly
to 11mm long with spreading or curved tips. Leaves affected the lower elevations on the islands of the
on saplings and young trees broader than on mature archipelago. Most collections date from before 1970,
trees, to 20mm wide, thin and flexible. Leaves of but recently it was collected from Choiseul and Santa
mature trees on distal parts of branches only, spread- Isabel. This species is not known from any protected
ing wide or more or less drooping, 1025cm long, areas.
712mm wide, linear-lanceolate, gradually tapering IUCN: EN [B2ab (iii)]
924 to a narrow petiolate, twisted base (leaving crescent-
shaped pulvini on the shoot when fallen) straight or Uses
slightly falcate, tapering to an acute or obtuse apex,
thick coriaceous. Midrib prominent on both sides This species, where it develops into a tree of mod-
of leaf, extending the full length, more or less acute erate size with a straight bole, is a useful timber
on adaxial (upper) side, 0.50.7mm wide, obtuse, tree producing soft, light wood suitable for light
becoming flattened distally and to 1mm wide on construction, interior finishing and boxes or cases.
abaxial side. Leaf colour lustrous green above, dull Locally it is being used to make oars. This species is
green below. Stomata small and inconspicuous, in not known to be in cultivation.
numerous intermittent lines on either side of abax-
ial midrib. Pollen cones unknown. Seed cones in
upper leaf axils on slender 1015mm long pedun- Podocarpus sellowii Klotzsch ex Endl., Syn. Conif.:
cles; receptacles subtended by 2 subulate, ca. 4mm 209. 1847.
long, acute bracts (foliola), obconical, slightly flat-
tened, 810mm long and broad, ca. 5mm thick at Etymology
apex, swelling to succulent red. Seeds 1, sometimes
2 per receptacle, including the epimatium subellip- This species was named after the German botanist
tical, 11 8mm, narrowed at base and rounded at Friedrich Sellow [Sello] (17891831).
apex, smooth and glaucous green. Seed proper not
observed. Vernacular names
Podocarpus spathoides de Laub., Blumea 30 (2): Podocarpus spathoides, first described and named
267. 1985. Type: Malaysia: Peninsular Malaysia, by De Laubenfels in 1985 (op. cit.), was based on his
Johore, Gunung Ledang [Mt. Ophir], D. J. de collection P 600 from Gunung Ledang [Mt. Ophir]
Laubenfels P 600 (holotype L). in Peninsular Malaysia. The isotype specimen at K
from a mature but small tree has elliptic to linear-
Etymology oblong, obtuse or obtuse-mucronate to apically
rounded leaves 58cm long and 1213mm wide,
The species epithet comes from Latin spatha = a with a raised midrib on the upperside of the leaf
broad flat wooden or metal blade and refers to the and a flat to shallowly grooved midrib on the lower
shape of the leaves. (abaxial) side. On that mountain it occurs from the
upper limit of P. ridleyi ( a species with lanceolate,
Vernacular names acute leaves) at ca. 1000m to the summit at 1276m.
It is a poorly known species based on very few
No common names are recorded for this species. collections.
Description Distribution
Small trees 36m tall, up to 20cm d.b.h. Bark not Malesia: Peninsular Malaysia (Gunung Ledang =
described. Branches spreading, forming a rounded Mt. Ophir), Sarawak (Lawas).
or domed crown. Foliage branchlets terete, more TDWG codes: 42 BOR-SR MLY-PM
Ecology Description
On Gunung Ledang [Mt. Ophir] this species occurs Shrubs to 3m tall, usually sprawling and not more
at altitudes from ca. 1000m to the summit at 1276m than 1.5m tall, layering. Bark thin, fibrous, brown.
a.s.l. in low shrubby vegetation and stunted forest. Foliage branchlets slender, terete, finely grooved, ter-
It is apparently rare there as on a visit in September minating in small, 24mm long buds with narrowly
2008 with staff from FRIM I did not find it in the triangular scales ending in fine, spreading apices.
summit area. Leaves on young and adult plants similar, narrowly
linear-lanceolate to linear, 2.57cm long, 24.5mm
Conservation wide, straight or slightly curved, short petiolate or
928 subsessile, gradually tapering to base and to an apicu-
The only certain locality of this species is the type late, pungent apex. Midrib acutely raised but narrow
locality, Gunung Ledang (Mt. Ophir) in Peninsular (0.3mm wide) on adaxial (upper) side, prominently
Malesia. Another possible locality is in Lawas, N raised and slightly wider on abaxial side. Leaf colour
Serawak, Borneo. The natural vegetation on Gunung yellowish green or lustrous green above, glaucous
Ledang has been influenced by development, involv- green below. Stomata in two conspicuous bands on
ing a road and television transmission towers and either side of abaxial midrib, forming intermittent
their infrastructure. It is apparently rare on this lines. Pollen cones axillary, solitary or in clusters of
mountain. If taken as the only locality known then 25 on 14mm long peduncles with minute scale
P. spathoides would be Endangered on the basis of leaves, short cylindrical, 48mm long, 1.52.5mm
its very restricted EOO in a partly disturbed moun- wide; microsporophylls minute, imbricate, finely
tain forest habitat. As the identity and status of the apiculate, bearing two basal, globose pollen sacs.
Sarawak occurrence are unknown but may belong Seed cones axillary to reduced leaves, full-grown
here, the actual status remains DD. leaves or subterminal, solitary on slender, 510mm
IUCN: DD long peduncles; receptacles subtended by two nar-
rowly lanceolate, curved or revolute 35mm long
Uses bracts (foliola), growing and swelling to 610mm
long, 67mm wide, succulent and purple when ripe.
No uses have been recorded of this species. Seeds obliquely attached to distal end of receptacle,
ovoid when mature, 810mm long, 57mm wide,
often distally crested; epimatium glaucous, becom-
Podocarpus spinulosus (Sm.) R. Br. ex Mirb., Mm. ing dark purple. Seed proper not observed.
Mus. Hist. Nat. 13: 75. 1825. Taxus spinulosa
Sm., in Rees, Cyclopaedia 35, Taxus No. 7. 1819; Distribution
Margbensonia spinulosa (Sm.) A. V. Bobrov &
Melikyan, Bjull. Moskovsk. Obsc. Isp. Prir., Otd. Australia: New South Wales, Queensland.
Biol. 103 (1): 60. 1998. Type: Australia: New South TDWG codes: 50 NSW-NS QLD-QU
Wales, Sydney Harbour, [Port Jackson], A. Philip
s.n. (holotype LINN). Fig. 289, 290, 291 Ecology
Huapsay, Guabisay (Quechua); romerillo (Spanish) Podocarpus sprucei is a high montane to subalpine
species growing in cloud forest up to the tree line, at
Description altitudes from 1800m to 3900m a.s.l. Only in forests
at lower altitudes does it become a tree to 20m tall.
Trees to 20m tall (dwarfed at highest altitudes);
trunk to 50cm d.b.h. Bark becoming scaly, reddish Conservation
brown. Branches ascending or spreading, forming
a bushy crown. Foliage branchlets slender, terete, Overexploitation for timber, logging the larger
finely grooved between leaf apices; terminating in trees, was observed in the field by the plant collector
clusters of small, verticillate buds or single, subglo- W. H. Camp (vicinity of Cuenca, Province of Azuay,
Ecuador) in 1945, resulting in scarcity of the resource patent, (3)48cm long, straight or slightly falcate,
in that area. Continuing exploitation is causing fur- (6)811mm wide, thick coriaceous, lanceolate to
ther decline of this species, which has an area of lanceolate-linear, narrowing to a petiolate, often
occupancy (AOO) of less than 500 km with a now twisted base, slightly V-shaped transversely or nearly
severely fragmented population. flat; margins slightly revolute, gradually tapering to
IUCN: EN [B2ab (ii, iii, v)] an acute or obtuse apex; leaf colour lustrous dark
green above, paler green below. Midrib on adaxial
Uses (upper) side slightly raised with a central groove, or
only a groove continuous to apex, obtusely raised
The wood of this species is much in demand and and 1mm wide on abaxial side. Stomata small, in
930 trees are logged from primary forest often unsus- numerous intermittent lines on either side of abaxial
tainably due to scarcity and slow growth. It is used midrib. Pollen cones axillary, solitary, sessile, sub-
in house construction and to make furniture as it tended by acuminate bud scales, cylindrical, length
takes a fine polish. This species is also planted as an at mature stage not observed. Seed cones axillary,
ornamental in parks in Ecuador. solitary on 718mm long peduncles; receptacles
composed of an axis with two unequal, fused and
slightly swollen bracts, 78mm long, becoming red
Podocarpus steyermarkii J. T. Buchholz & or purple. Seeds including the epimatium ovoid-glo-
N. E. Gray, J. Arnold Arbor. 29: 133. 1948. Type: bose, ca. 8mm long, with a minute crest.
Venezuela: Bolvar, Rio Carrao, Carrao-tepui,
J. A. Steyermark 60876 (holotype F). Pl. 39 Distribution
Plate 39. Podocarpus steyermarkii. 1. Branch with leaves and seed cones. 2. Seed cone. 3. Branch with
leaves and pollen cones. 4. Microsporophyll (dried). 5. Foliage bud and leaves. 6. Bud scale. 7. Cross-section
of leaf.
Conservation adaxially, wider and centrally grooved abaxially. Leaf
colour bright green above (adaxially), pale green
This species, which was described and known from below; new leaves flushing yellowish green. Stomata
only a single locality in 1948, is still known from only on abaxial (under) side, very small, in numerous
six or seven localities. It is likely to turn up in more irregular lines on either side of midrib. Pollen cones
places with continuing botanical surveys, but is axillary, sessile, in clusters of (1)25, subtended
obviously scattered and uncommon. Its total area of by scarious, rostrate, keeled scales 35mm long,
occupancy (AOO) is almost certainly less than 500 cylindrical; cones elongating to 2040(60?)mm
km2, but it is not exploited, nor are the forest patches 2.53.5mm at anthesis; microsporophylls spirally
in which it occurs threatened by deforestation. This arranged, with a triangular, spreading, 0.3mm long
932 species is therefore considered to be outside any cat- apex and with two pollen sacs. Seed cones solitary
egory of threat of extinction in the wild. and axillary on 515mm long, slender peduncles;
IUCN: LC receptacles 712mm long, with two 12mm long
bracts (foliola) at their base, swelling when ripe to
Uses a succulent, red pseudo-fruit ca. 10mm thick. Seeds
single (sometimes 2) at truncate distal end of recep-
No uses have been recorded of this species and it is tacle, obliquely globose, 1012 810mm including
not known to be in cultivation. the green epimatium, minutely crested distally, rip-
ening to blackish purple. Seed proper ca. 8 7mm,
slightly flattened, brown.
Podocarpus subtropicalis de Laub., Blumea 30 (2):
277. 1985. Type: China: Sichuan, Emei Shan, Taxonomic notes
[Mt. Omei], E. H. Wilson 3007 (holotype A).
Podocarpus subtropicalis has been identified with P.
Podocarpus subtropicalis de Laub. var. medogensis neriifolius by Chinese botanists (see Flora of China
Silba, J. Int. Conifer Preserv. Soc. 7 (1): 40. 2000. 4: 82, 1999), although it is usually not explicitly syn-
onymized with that species in the recent floristic
Etymology literature. Its taxonomic status remains somewhat
uncertain and is not helped by the circumstance that
The species epithet refers to the subtropical habitat it seems to have been widely spread in cultivation.
or distribution of this species. Its morphology is also close to P. fasciculus, a species
from Taiwan described by De Laubenfels (1985) in
Vernacular names the same paper on the same page, both with short
Latin descriptions and type designations, but with-
No vernacular name is recorded for this species. out comparative discussion. I have retained these
species here as distinct, but more critical taxonomic
Description comparison based on further sampling from trees
occurring in the wild is needed.
Trees to 20m tall, monopodial. Bark not described
or observed. Foliage branches terete, grooved, termi- Distribution
nating in buds of 45(7?) 23mm, with lanceo-
late, keeled and spreading or recurved scales. Leaves China: Sichuan (Emei Shan [Mt. Omei]), Yunnan.
linear-lanceolate, petiolate (petioles 37mm long), TDWG codes: 36 CHC-SC CHC-YN
on juvenile plants up to 18cm long and 14mm wide;
on adult plants 512(15)cm long, (5)713mm Ecology
wide, coriaceous, stiff, straight or more or less fal-
cate, gradually tapering at both ends, acute or acumi- This species was described and named in 1985 by
nate. Midrib prominent on both sides, acutely raised David J. de Laubenfels, based on a collection from the
famous mountain with Buddhist shrines and stair- or less tesselated, brown. Branches on trees spread-
way paths Emei Shan [Mt. Omei] in Sichuan, China. ing to form a broad crown, densely set on shrubby
Little is known of the ecological circumstances and trees, spreading out wide. Foliage branchlets terete,
many trees have been introduced there over the cen- finely grooved, terminating in small, globose or
turies. It was subsequently reported from Yunnan, ovoid buds with 12mm long, imbricate, appressed
but it remains a poorly known taxon. It is also widely and obtuse scales. Leaves on juvenile plants larger
planted elsewhere in China and, according to David than on mature plants, to 16cm long and 18mm
de Laubenfels, in many other parts of the world with wide. Leaves on mature trees and shrubs variable,
a warm-temperate to subtropical climate. (3)510(13)cm long, (5)711(13)mm wide, oval-
linear (widest at midpoint) to linear, tapering grad-
Conservation ually down to a more or less petiolate base and to 933
an obtuse apex, coriaceous, straight or only slightly
IUCN: DD curved. Midrib prominent but obtusely raised on
adaxial (upper) side, 11.2mm wide, flat or forming
Uses a groove in dried leaves on abaxial side. Leaf colour
bright lustrous green above, dull light green below.
This species, occurring on Emei Shan [Mt. Omei], Stomata very small, in two bands forming numer-
has been taken from there and seeded and/or planted ous intermittent lines on abaxial side. Pollen cones
widely in other parts of China and beyond. It is often from clustered buds in axils of leaves or below
identified as P. neriifolius. these, sessile, solitary or 23 together, subtended by
small, rounded, imbricate bud scales, cylindrical,
1020mm long, 3mm wide, yellow turning brown;
Podocarpus sylvestris J. T. Buchholz, Bull. Mus. microsporophylls imbricate, peltate, with erose-
Hist. Nat. (Paris), sr. 2, 21: 285. 1949. Type: New denticulate margin and minutely apiculate, bearing
Caledonia: Grande Terre, Province Sud, Fort du two basal, globose pollen sacs. Seed cones axillary
Mois de Mai, J. T. Buchholz 1696 (holotype ILL). and solitary on slender, 715mm long peduncles;
receptacle subtended by two small bracts (foliola),
Podocarpus novae-caledoniae Vieill. var. colliculatus growing to 610mm long, swelling to 56mm
N. E. Gray, J. Arnold Arbor. 39: 432. 1958; Podocarpus thick, maturing to red or red-brown. Seeds includ-
colliculatus (N. E. Gray) de Laub., Adansonia, ser. 3, ing the epimatium relatively large, 1015 79mm,
27 (2): 152. 2005. obliquely ovoid with a faint crest, smooth, olive
green turning brown. Seed proper ovoid, smooth,
Etymology ca. 9 6mm.
The species epithet sylvestris means from the forest Taxonomic notes
and refers to its habitat.
De Laubenfels (op. cit.) elevated var. colliculatus
Vernacular names of N. E. Gray (op. cit.), which she had placed with
Podocarpus novae-caledoniae and described from
No common names have been recorded of this the le des Pins in New Caledonia, to a species. Gray
species. should have placed this taxon (as a variety) with P.
sylvestris, from which it differs only slightly in hav-
Description ing somewhat larger receptacles and longer pedun-
cles. Both P. novae-caledoniae and P. sylvestris occur
Small to medium size trees to 20m, rarely to 30m on the le des Pins and are well distinguished mor-
tall and 80cm d.b.h., or at high altitude shrubby trees phologically, but the differences between P. collicula-
1.53m tall. Bark on large trees thick and fibrous, fis- tus and P. sylvestris are minor and not constant in the
sured longitudinally, or on small trees thin and more specimens I have seen.
Distribution Podocarpus tepuiensis J. T. Buchholz & N. E. Gray,
J. Arnold Arbor. 29: 134. 1948. Type: Venezuela:
New Caledonia (mountainous parts of Grande Bolivar, La Gran Sabana, Ptari-tepui, J. A.
Terre, le des Pins). Steyermark 59716 (holotype F).
TDWG codes: 60 NWC
Etymology
Ecology
The species epithet refers to the (region of the) tepuis
Podocarpus sylvestris grows as a scattered tree in or table mountains of Venezuela.
the tropical evergreen rainforests that cover parts of
934 the mountains of New Caledonia; it is also found Vernacular names
in wooded ravines and at high altitude in vegeta-
tion bordering on maquis minier, the shrubby veg- No common names have been recorded for this
etation on serpentine. It is not restricted to these species.
ultramafic soils, as it also occurs on micaschist in
the far NE of the main island Grande Terre. The alti- Description
tudinal range is 150m to 1200m a.s.l. At high alti-
tude or on ultramafic rocks it becomes a shrub or Shrubs or small trees 16m, rarely to 15m tall;
shrubby tree of low stature; in well developed rain trunks on the largest trees to 30cm d.b.h. Bark
forest it can reach 2030m tall. Besides numerous thin, fibrous, reddish brown. Branches ascending,
angiosperms, its associated species are sometimes forming a (narrowly) pyramidal, dense crown in
conifers, mainly Araucaria spp. and Agathis spp. It small trees, spreading more horizontally in shaded
is most often found as a small understory tree with trees and with more or less erect foliage in shrubs.
these conifers, but is fertile in these circumstances Foliage branchlets slender, terete, with fine grooves
just as in more open maquis-like vegetation or as a between leaf bases, terminating in small, globose
canopy tree. buds 23mm long and wide, with inbricate, broadly
triangular, carinate scales with narrow scarious mar-
Conservation gins. Leaves densely set, spreading at wide angles to
shoot; on juvenile plants up to 6cm long and 8mm
This species has been the target of logging, an activ- wide; on mature plants (1)1.52.5(3.5)cm long,
ity which has undoubtedly led to a reduction in its oblanceolate or elliptic, 3.55mm wide, often slightly
abundance, but is difficult to quantify. Such opera- or markedly curved down, gradually narrowing
tions have now virtually ceased. This species is to a short petiolate base; margins slightly revolute;
widely distributed and still locally common and if apex obtuse or more or less acute; leaf texture coria-
forests are allowed to regenerate it will again spread ceous; leaf colour deep lustrous green above, pale
from these localities. green below; young leaves flushing pinkish white.
IUCN: LC Midrib on adaxial (upper) side of leaf a continuous
groove, inconspicuous and nearly flat on abaxial
Uses side. Stomata very small, in numerous intermittent
lines on abaxial face of leaf. Pollen cones axillary,
This species has been subject to logging in the past; solitary, sessile with rounded perular scales at base,
its wood provides good material for construction cylindrical, 1015mm long, 1.5mm wide, cream
and indoor carpentry, floor boards, and furniture coloured when mature; microsporophylls triangular,
and has been known as false kauri in the timber with scarious margins, bearing two globose pollen
trade. Exploitation has virtually ceased as the more sacs at base. Seed cones axillary, solitary on a short
accessible forests have been logged and forest con- (23mm long) peduncle; receptacles constituting
servation of the remainder is now the prevailing of an axis with two unequal, divergent but mostly
management practice. This species has been planted fused, slightly swollen, fleshy bracts, the largest fer-
in some parks and gardens in New Caledonia. tile, ca. 5mm long, green pruinose ripening to red
or purple. Seeds including the epimatium 67mm Uses
long, 4mm wide, obliquely ovoid, with a distal crest
forming a papillate protrusion. No uses have been recorded of this species and it is
unknown in cultivation.
Distribution
Ecuador: Cordillera del Condor, Cordillera Sacha Podocarpus teysmannii Miq., Fl. Ned. Ind. [Fl. Ind.
Llanganates; Venezuela: Bolivar, Amazonas; Guyana: Batavae] 2 (7): 1072. 1859. Podocarpus neriifolius D.
Cuyuni-Mazaruni Region. Don var. teysmannii (Miq.) Wasscher, Blumea 4 (3):
TDWG codes: 82 GUY VEN 83 ECU 453. 1941. Type: Indonesia: Sumatera, Utara, on the
coast near Sibolga, J. E. Teijsmann s.n. (holotype L). 935
Ecology
Etymology
Podocarpus tepuiensis occurs in the mountains
(tepuis = table mountains and other mountains) as This species epithet commemorates the Dutch bota-
well as in the lowlands. The altitudinal range in the nist Johannes Elias Teijsmann (18091882).
mountains is from 700m to 2450m a.s.l.; the two
herbarium collections in the southern Venezuelan Vernacular names
lowlands are from 100110m a.s.l. In the latter local-
ity it occurs on the edge of forest near savanna, on On the west coast of Sumatera this species is known
white sand with seasonal flooding (black water), and as kalek rotan; Pilger (1903) cited the name Sikuju
with occasional fires on the savanna; associated spe- laut, presumably also from Sumatera.
cies are e.g. Henriquezia nitida, Leopoldina piassaba,
Emmotum sp., and Aldina sp. On the Venezuelan Description
tepuis it occurs on slopes and summits, usually
along streams in gallery forest or at higher altitudes Trees to 36m tall, commonly to 20m, d.b.h. to
in dwarf forest patches. It may sometimes be associ- 50cm; trunk erect, straight, terete; large trees with
ated with P. roraimae and P. steyermarkii, but these slightly fluted base. Bark smooth, eventually with
two podocarps do not occur at lower altitudes than small, powdery scales, brown; inner bark soft and
1800m a.s.l. and in most localities only one of these fibrous, orange-brown. Branches spreading, forming
species has been found. The disjunct localities in a rounded crown. Foliage branchlets terete, slightly
Ecuador, where this species has been found since grooved or ridged, terminating in globose compact
1993 are at altitudes between 1600m and 2740m. It buds of 36mm diam. with imbricate, rounded
occurs on quarzite outcrops in a sandstone forma- scales with erose margins. Leaves on juvenile plants
tion similar to the Guyana Shield mesa sandstones lanceolate, to 20cm long and 22mm wide, straight
and on granite. or falcate, thin, gradually tapering to an acute apex.
Leaves on mature plants lanceolate to linear-lan-
Conservation ceolate, 714cm long, (10)1217(20)mm wide,
straight or slightle curved, thinly coriaceous, grad-
Podocarpus tepuiensis is the most common and ually tapering to a petiolate base; apex acute or
widespread of the tepuis podocarps and actually acuminate. Midrib on adaxial side narrow, acutely
occurs elsewhere, too. It is a shrub or small tree of raised from base to apex, on abaxial side wider, to
little interest for timber and most of the popula- 1.5mm, obtusely raised. Stomata very small, in
tions are remote from roads and towns. The disjunct numerous irregular lines on abaxial (under) side.
occurrence in Ecuador makes it likely that its total Pollen cones axillary, solitary or in clusters of 23,
range is incompletely known at present. sessile, subtended by rounded bracts (bud scales),
IUCN: LC narrowly cylindrical, 2540mm long, ca. 3mm
wide; microsporophylls spirally arranged, spreading, Podocarpus totara G. Benn. ex D. Don, in Lambert,
with triangular apex and two basal pollen sacs. Seed Descr. Pinus, ed. 2: 184. 1828. [& in ed. 8, 2: [189].
cones axillary, solitary on a 310mm long, slender 1832]. Type: New Zealand: North Island, Bay of
peduncle; receptacles subtended by 2minute, early Islands, [?] Bennett s.n. (lectotype BM, [specimen a]
deciduous bracts (foliola), slender, ca. 10mm long, designated here). Fig. 293, 294, 295
consisting of two lobes (axes?) each with a dis-
tal bract, one of these being fertile; lobes swelling Podocarpus totara G. Benn. ex D. Don var. waihoen-
mostly distally, ripening to purplish pruinose. Seeds sis Wardle, New Zealand J. Bot. 10 (1): 201. 1972.
single, obliquely attached, including the epimatium
79 57mm, subglobose with a small crest, dark Etymology
936 green turning blackish brown.
The species epithet uses the Maori name for this tree.
Distribution
Vernacular names
Malesia: Peninsular Malaysia, Sumatera (including
Bangka & Lingga Islands), Borneo (Brunei, Sabah). totara (Maori)
TDWG codes: 42 BOR-BR BOR-SB MLY-PM SUM
Description
Ecology
Trees to 30(40) m tall, with a straight, sometimes
In Flora Malesiana, ser. 1, 10 (3): 407 (1988) massive trunk to 3.6m d.b.h. Bark thick, fibrous,
Podocarpus teysmannii was described as an under- stringy, with longitudinal furrows, exfoliating
storey tree to 12m tall, but in fact sometimes trees in small, fragmentary strips, light grey-brown.
are found reaching the canopy with a height of 36m Branches spreading, forming a rounded crown, in
(T. C. Whitmore FRI 8835collected on Gunung veteran trees irregular through much reiteration.
Blumut in Peninsular Malaysia at 510m on a ridge). Foliage branchlets terete, glabrous, finely grooved;
This species occurs in both primary and secondary branching irregularly, sometimes dense, but shaded
rain forest from near sea level to occasionally 1350m branchlets more or less pectinate. Terminal buds
a.s.l., but mostly up to 800m. It is often restricted to 23mm long, subglobose, the inner scales imbri-
sites with poor, sandy, or rocky soils and therefore cate and obtuse, the outer scales acute with free
will not attain large size under these conditions. On tips. Leaves sessile, short decurrent, spreading at
the coast of Sabah it often occurs on sandy ridges wide angles, lanceolate-linear, (0.5)12.5(3) cm
or hills with the broadleaved tree Dryobalanops long, (1.5)23.5(4.5)mm wide (narrowest leaves
rappa (Dipterocarpaceae); in the Mt. Kinabalu area on saplings and young trees, widest leaves on fertile
(Mesilau River) it was collected in kerangas forest branches of large trees), straight or slightly curved,
dominated by Agathis sp. tapering to a slightly twisted base and an acute-
pungent or sometimes obtuse-mucronate apex,
Conservation coriaceous, stiff; a faint groove through the middle
adaxially (upperside) and an obtuse midrib abaxi-
IUCN: NT ally; leaf colour dark green, more or less lustrous
above, dull and lighter green below. Stomata in two
Uses irregular bands of intermittent lines on abaxial side.
Pollen cones axillary, solitary or sometimes in pairs
Larger trees of this species are undoubtedly logged on short peduncles, subtended by small, triangular
for their timber; the use of which will be similar to scales, cylindrical, 1215(20)mm long, 34mm
that of other podocarps of the region. Podocarp wide at anthesis, straight or curved after shedding
wood is suitable for light construction, carpentry pollen, yellowish green turning brown; microspo-
and joinery, furniture making, etc. rophylls imbricate, peltate-triangular, upper margin
denticulate, with two sublateral, globose, yellow ally it penetrates into this zone at around 500m a.s.l.
pollen sacs. Seed cones axillary, solitary on a short Above this altitude it is replaced by the similar spe-
peduncle; receptacles when growing narrowly cies P. cunninghamii.
oblong, with two distal bracts subtending 12 ovules,
swelling to nearly globose, 4.55.5mm long, yellow Conservation
ripening to scarlet, soft and succulent. Seeds includ-
ing the epimatium ovoid-oblong, with a constricted Under the A criterion of IUCNs Red List of
distal part, 33.5 22.5mm, lustrous green or glau- Threatened Species, this species would have to
cous green. Seed proper ca. 2.5mm long, narrowly be listed in a threatened category due to relatively
ovoid; seed coat smooth and hard. recent major decline, caused by logging and forest
clearance in the past 150 years. In New Zealand, this 937
Taxonomic notes criterion, if applied to native trees, is not accepted by
conservationists, due to more recent legislation that
A herbarium sheet of this species at BM contains a gives protection to most of the remaining forests and
mixture of specimens collected by different people at trees, both within reserves and through legal bans on
different localities and times. The largest branchlet cutting and other means of clearance of native forest.
marked a) was collected by Bennet in July 1829 at the This policy has in fact already led to an increase in
Bay of Islands. Brian Molloy annotated this as the abundance (from an all-time low just before forest
lectotype of Podocarpus totara in October 1991, but clearance was prohibited), due to regeneration and
since he never published this, that lectotypification spread from seed trees into secondary vegetation
is effected here. and pioneer phases of native forest. As demand for
further conversion of forested land for agriculture
Distribution (diary, sheep farming) is presently waning, for the
foreseeable future Podocarpus totara and other low-
New Zealand: North Island, South Island. land conifers are indeed out of danger. However, this
TDWG codes: 51 NZN NZS situation is policy dependent and there is no abso-
lute guarantee that this policy will never change in
Ecology the other direction in the future. The current situa-
tion indeed does not warrant listing of this species in
Podocarpus totara was the largest conifer dominat- a threatened category.
ing the lowland forests that covered much of the two IUCN: LC
large islands of New Zealand before humans, and
especially Europeans, intervened. It now occurs only Uses
in forest fragments, often in secondary vegetation.
It is a slow growing, emergent tree dependent in The Maori have used totara (not exclusively P.
that ecosystem upon episodal events of disturbance totara) for many purposes, but especially construc-
enabling massive regeneration. Past forest clearance, tion of houses. Europeans began large scale logging
if it took place in what are now forest rerserves, and forest clearing in the 19th century and much
can provide these conditions. Maximum age is ca. of the wood of totara went into fence posts on the
8001000 years and one tree is estimated to exceed establishing farms and later in that century for rail-
1800 years. In primary forest this species is often way sleepers. Only the best trees were sawn in the
associated with other conifers, e.g. Dacrycarpus numerous sawmills that had sprung up everywhere,
dacrydioides, Dacrydium cupressinum, Prumnopitys the remainder were wasted (Ogden & Stewart
ferruginea, Phyllocladus trichomanoides, and in in Enright & Hill, 1995). Timber export between
Northland Agathis australis. Angiosperm trees 18501900 included totara, but its main source was
can also be common and are often varied. Usually kauri (Agathis australis) until that resource was
Podocarpus totara grows well below the altitude almost exhausted. The wood is reddish, but vari-
where Nothofagus begins to dominate, but occasion- able in hue, dense and straight-grained, and was
used for general construction, joinery, furniture 5(6.5)cm long, (3.5)47(9)mm wide, narrowly
and cabinet making, as well as the coarser appli- lanceolate to oblanceolate, straight, gradually nar-
cations like railway sleepers, posts for communi- rowing to a short petiolate base, gradually or more
cation lines, and fencing. At present, native trees abruptly tapering to an acute or obtuse apex; mar-
are virtually universally protected in New Zealand gins not parallel for most of leaf length, straight
and the exploitation of the wood of totara is much or slightly revolute; leaf texture coriaceous or thin
restricted, essentially to dead trees outside reserves, coriaceous; leaf colour green or grey-green above,
and put to specialist wood work only. In horticul- whitish green below. Midrib 0.50.7mm wide, con-
ture it remains uncommon, mostly restricted to tinuous or sometimes fading towards apex, mostly
arboreta and botanic gardens. Although eventually forming a groove on adaxial (upper) side, more or
938 growing to a large tree if left alone, it can be planted less raised from a flat leaf surface and continuous
and trimmed for hedges. to apex on abaxial side. Stomata small, in numer-
ous intermittent lines forming two bands on abaxial
side. Pollen cones axillary, on short shoots or slen-
Podocarpus transiens (Pilg.) de Laub. ex Silba, der peduncles up to 15mm long in 1-several clus-
Phytologia Mem. 7: 68 (1984). Podocarpus ters of 26 or more, subtended by apiculate perular
lambertii Klotzsch ex Endl. var. transiens Pilg., scales, cylindrical, 612mm long, 1.52mm wide;
in Engler, Pflanzenr. IV.5 [18]: 86. 1903. Types: microsporophylls ovate-triangular, with minutely
S Brazil, F. Sello s.n.; A. F. M. Glaziou 16335 denticulate margins, bearing two globose pol-
(syntypes, not located). len sacs. Seed cones axillary, solitary on slender,
510mm long peduncles; receptacles small, swell-
Podocarpus transiens (Pilg.) de Laub. ex Silba var. ing to 67mm long, 45mm wide, succulent,
harleyi Silba, J. Int. Conifer Preserv. Soc. 7 (1): 39. purple, eventually chestnut brown. Seeds solitary,
2000. globose, including the epimatium 45 45mm,
glossy without a crest. Seed proper globose with a
Etymology smooth surface.
This species is an uncommon component of mon- No common names have been recorded for this
tane forest remnants surrounded by campo rupestre species.
(rocky grassland), a dryland vegetation in the inte-
rior. Nearer the Atlantic coast and in the south of its Description
range it has been found in remnants of Atlantic Rain
Forest. The altitudinal range of this species, based on Trees to 1520m tall; trunk with d.b.h. to 6080cm,
data on herbarium specimen labels, is 10001780m often fluted at base. Bark smooth, on large trunks
a.s.l. It may be associated with Podocarpus lamber- exfoliating in small strips, dark brown weathering
tii, especially in the southern parts of its range, but grey. Branches spreading, forming a wide crown 939
unequivocal evidence of this has not been recorded in old trees. Foliage branches terete, finely grooved
on herbarium labels or in the literature, probably between leaf bases, terminating in small subglobose
due to identification problems, as most specimens buds with free, triangular, apiculate outer scales and
were identified as P. lambertii. imbricate inner scales. Leaves on shaded branches
and juvenile trees larger than on sun-exposed
Conservation branches of mature trees, 610cm long, 1015mm
wide, straight or sometimes slightly falcate. Leaves
Due to very small numbers of trees in some of its on sun-exposed branchlets (2.5)46(7)cm long,
locations, the area of occupancy (AOO) must be 612mm wide, spreading or directed forward on
much smaller than 500 km2, while populations ends of shoots, coriaceous, dark green above, pale
appear always to be small in the ca. 10 locations green below, straight, elliptic to oblanceolate or
recorded in the region, of which the most com- oblong, gradually narrowing to a short petiolate base
monly visited is the Morro do Chapeu in Bahia. In and more abruptly to an acute to weakly apiculate or
several locations degradation of habitat and tree cut- rarely obtuse apex. Midrib on adaxial (upper) side
ting for firewood are causing decline of this species. prominently raised but sometimes fading towards
Deforestation caused by cutting, fire and grazing of apex, less than 0.5mm wide and lying in a shallow
domestic animals is the main threat. The tree cutting depression, on abaxial side flat or forming a shallow
is mostly for firewood, much less for timber. groove (in sicco). Stomata small, in numerous inter-
IUCN: EN [B2ab (ii, iii, v)] mittent, irregular lines on abaxial side. Pollen cones
axillary, solitary, sessile, subtended by thin, scarious
Uses bud scales, cylindrical, maximum elongation not
observed; microsporophylls triangular, minutely
No commercial uses have been recorded for this apiculate, bearing two relatively large, ovoid-oblong
small tree, of which the timber has not much value. pollen sacs. Seed cones axillary on short peduncles;
It is used locally for fence posts and more extensively basal bracts (foliola) deciduous; receptacles formed
for firewood. It is not known to be in cultivation. of an axis with two unequal, fused bracts, swelling to
a 78 6mm large, succulent, red body. Seeds soli-
tary, including the epimatium ovoid, 78 56mm,
Podocarpus trinitensis J. T. Buchholz & N. E. Gray, with or without a distal, curved, thin crest. Seed
J. Arnold Arbor. 29: 135. 1948. Type: Trinidad and proper ovoid, smooth with a thin, light brown seed
Tobago: Trinidad, Saint Andrew Co., Valencia, coat.
[Valencia road], W. E. Broadway 7151 (holotype
MO). Taxonomic notes
Plate 40. Podocarpus urbanii. 1, 2. Branches with leaves and seed cones. 3. Branchlet with leaves and
pollen cones. 4. Leaves. 5. Seed cone.
Ecology status in the IUCN Red List 2008. It has a limited
distribution confined to a single mountain range,
Podocarpus urbanii occurs in montane to high mon- where it is locally common. A recent re-assessment
tane forests at altitudes between 1160m and 2256m in 2011 has found that it is Critically Endangered due
a.s.l. in the Blue Mountains of E Jamaica. to its limited occurrence and continuing decline.
IUCN: CR [B1ab (iiii, v)]
Conservation
Uses
In the Global Red List of Conifers (Farjon & Page,
1999) this species was listed as Near Threatened No economic uses have been recorded of this
942 (LRnt following 1994criteria) and this was still its species.
Prumnopitys Phil., Linnaea 30: 731. 1861. Type: Prumnopitys andina (Poepp. ex
Endl.) de Laub. (Podocarpus andinus Poepp. ex Endl.) [Prumnopitys elegans Phil.]
(Podocarpaceae).
Stachycarpus Tiegh., Bull. Soc. Bot. France 38: 163. observable in the field are insufficient to differenti-
1891. Type: Stachycarpus andinus (Poepp. ex Endl.) ate between them. Therefore pollen cones and ripe
Tiegh. [Prumnopitys andina (Poepp. ex Endl.) de seeds are necessary to arrive at a correct identifica-
Laub. (Podocarpus andinus Poepp. ex Endl.)] tion if one has to make a choice from among all nine
species.
Greek: prumme, prumne = stern; pitys = pine or fir 943
tree. 1a. Adult leaves spreading to all sides (but more or
less pectinate on shaded shoots), slightly
Description curved backward or assurgent, slender,
(1)1.52mm wide P. taxifolia
Trees or multistemmed shrubs, evergreen, dioecious, 1b. Adult leaves pectinately arranged, nearly
with smooth or scaly bark. Resin canals in leaves (1) straight or slightly curved sideways, sometimes
and resin in bark. Foliage branches alternate, spread- falcate, (1.8)23.8mm wide 2
ing in a more or less horizontal plane. Leaves small, 2a. Small trees to 15 m tall, with a short bole or
spirally inserted, in vegetative lateral shoots usually multi-stemmed. Adult leaves 1.82.5mm wide.
pectinately arranged, flattened, more or less falcate- Seeds including the fleshy epimatium 1825
linear, epistomatic with stomata in two bands sepa- 1820 mm, bluish green ripening to yellow
rated by a thin midrib. Pollen cones aggregate on P. andina
relatively long, leafless branchlets, cylindrical, slen- 2b. Trees usually with a single bole and potentially
der; pollen sacs relatively large, yellow, two on each taller than 15m. Adult leaves 23.8mm wide.
microsporophyll, transversely dehiscent, containing Seeds including the fleshy epimatium 1020
bisaccate pollen. Seed cones on small axillary branch- 715mm, variably coloured (if larger than 20
lets with a few reduced leaves, consisting of one to a 15mm, dark blue to blue-black) 3
few spirally arranged bracts developing a single erect 3a. Adaxial (upper) midrib (midvein) of leaves dis-
ovule (seed) in the axil of a small (terminal) bract, tinctly grooved. Pollen cones arranged in axil-
lacking any fleshy receptacle at maturity. Seeds ovoid, lary spikes of 1525 4
with a drupe-like, thick, fleshy and globose or ovoid- 3b. Adaxial midrib of leaves indistinctly raised
elliptic epimatium (outer layer) covering the seed (not a groove). Pollen cones solitary, nearly
entirely and becoming yellow, red or black. sessile 7
4a. Leaf apex obtuse, sometimes acute or acumi-
9 species nate. Seeds including the fleshy epimatium
crested, pruinose when ripe 5
Distribution 4b. Leaf apex acute or acuminate, sometimes
mucronate or obtuse. Seeds including the fleshy
Australia: Queensland. SW Pacific: New Caledonia; epimatium rounded or with an apical knob,
New Zealand. America: Costa Rica (Central yellow or bluish, but not pruinose when ripe
America); in South America along the Andes from 6
W Venezuela to Bolivia; disjunct in S Chile. 5a. Seeds (cones) on 0.51.5 cm long branchlets
with deciduous scale leaves. Microsporophylls
Key to the species of Prumnopitys of pollen cones with an acute apex, 0.5 mm
wide P. exigua
The vegetative characters are often very simi- 5b. Seeds (cones) on 1.52.5 cm long branchlets
lar among the species of Prumnopitys and the few with persistent scale leaves at base and
different character states in the leaves that are easily deciduous reduced leaves. Microsporophylls of
pollen cones with broadly triangular apex, Vernacular names
0.7mm wide P. harmsiana
6a. Seeds (cones) on 0.51.5 cm long branchlets Plumyew; Lleuque, Lleuqui, Uva de Cordillera
with a few reduced leaves, solitary. Seeds (Chile)
including the fleshy epimatium 10 78mm,
without an apical crest or knob P. standleyi Description
6b. Seeds (cones) on 23cm long branchlets with
(near) normal leaves, solitary or in pairs. Seeds Trees to 15m tall; trunk to 1m d.b.h., with a short
including the fleshy epimatium 1416 bole or multi-stemmed. First order branches spread-
1012mm, with a distinct apical knob ing or ascending forming a broadly pyramidal or
944 P. montana rounded crown. Bark smooth on younger trees,
7a. Pollen cones very small, 34 1.31.6 mm. becoming scaly and exfoliating in irregular small
Seeds including the fleshy epimatium 1014 strips; bark colour purplish grey, weathering grey.
710mm, with a ridge and apical knob New foliage glaucous green, turning greyish green.
P. ferruginoides Seedlings and young plants with slender leaves
7b. Pollen cones larger, 720 23 mm. Seeds 815(20)mm long and 1.22mm wide; apex acu-
including the fleshy epimatium 1530 minate. Leaves on mature trees alternate, twisted
925mm, only with an apical knob 8 at a petiolate base, mostly pectinately arranged in
8a. Adult leaves 23mm wide, curved downward. two rows and one plane, linear, (8)1220(25)mm
Pollen cones 820 mm long. Seeds including long, 1.82.5mm wide, slightly curved sideways
the fleshy epimatium 1518 913mm, yellow above base or more or less falcate, with parallel
becoming bright red P. ferruginea sides, with a indistinct proximal midrib adaxially
8b. Adult leaves 2.53.5mm wide, (nearly) straight. (upperside) and a low midrib abaxially (under-
Pollen cones 79mm long. Seeds including the side); margins entire; apex acuminate or sometimes
fleshy epimatium 2230 1825 mm, dark obtuse; leaf colour dull greyish green above, whit-
blue to blue-black P. ladei ish green below. Stomata in two indistinct bands
of numerous lines, more or less separated by mid-
rib on abaxial (under) side of leaves. Pollen cones
Prumnopitys andina (Poepp. ex Endl.) de Laub., on lateral and axillary spikes towards ends of foli-
Blumea 24 (1): 189. 1978. Podocarpus andinus age branches; up to 25 spreading cones per spike,
Poepp. ex Endl., Syn. Conif.: 219. 1847. Type: 47mm long and 2mm wide, each subtented by a
[Habitat in Chile australis alpinis frigidis, locis leaf-like bract 3mm long. Microsporophylls sub-
umbrosis convallis Quillai-Leuvu versus Antuco. peltate with acute apex, 0.5mm wide, incurved at
Pppig msc.] (HAL?, W destroyed). Fig. 296, 297 anthesis, light green, each with two yellow pollen
sacs. Seed cones 13 on 23cm long branchlets
Podocarpus spicatus Poepp., in Poeppig, Nov. Gen. with persistent scale leaves at base and deciduous,
Sp. Pl. 3: 18. 1845, non R. Br. (1838); Prumnopitys acute scale leaves, axillary to a scale leaf (bract),
spicata (Poepp.) Molloy & Muoz-Schick, New Zeal. consisting of an epimatium enclosing a subglobose,
J. Bot. 37: 190. 1999 (nom. illeg., Art. 53.3). slightly flattened, smooth seed, growing into a glo-
Prumnopitys elegans Phil., Linnaea 30: 732. 1860. bose to broadly elliptical, bluish green ripening to
Prumnopitys andina (Poepp. ex Endl.) de Laub. yellow pseudo-fruit 1825 1820mm diam. with
subsp. blijdensteinii Silba, J. Int. Conifer Preserv. an apical small knob.
Soc. 9 (1): 33. 2002.
Distribution
Etymology
S Argentina: Neuqun; S Chile: Biobio, La Araucana,
The species epithet refers to its occurrence in the Maule.
Andes. TDWG codes: 85 AGS-NE CLC-BI CLC-LA CLC-MA
Ecology from earlier introductions. Growing anew from seed
would increase genetic diversity, thought to be cru-
Prumnopitys andina occurs in valleys close to riv- cial to counter potential pathogens.
ers, in mixed forests with Austrocedrus chilensis,
Cryptocarya alba, Quillaja saponaria and other
sclerophyllous shrubs and trees. In the Andes Prumnopitys exigua de Laub. ex Silba, Phytologia
and Coastal Cordillera it grows with Nothofagus Mem. 7: 68. 1984. Type: Bolivia: Cochabamba,
dombeyi, N. nervosa and N. obliqua in non-sclero- road from Sihuenca to Totora, M. Cardenas 4879
phyllous montane rain forests. Its altitudinal range is (holotype US).
from 200m to 1400m a.s.l.
Etymology 945
Conservation
The species epithet is from Latin: exiguus = short,
There are probably fewer than 12 subpopulations insignificant; this species was at first believed to be a
and only two of these have more than 1000mature stunted form, but larger trees were found later.
individual trees, while many have fewer than 100.
Several subpopulations have been reduced recently, Vernacular names
e.g. by conversion of the natural forest to plantations
of exotic trees for commercial forestry. Other threats pino colorado, pino negro (Spanish)
are clearing of woodland to make space for agricul-
ture, flooding in relation to hydroelectric projects, Description
and increased wildfires. Regeneration is high in
some subpopulations but very low in others, possi- Trees to 20m tall; trunk to 1m d.b.h., with a short,
bly due to domestic animals eating the fleshy seeds. stout bole. First order branches spreading in large
There are three protected areas in Chile which con- trees forming a rounded crown. Bark smooth on
tain this species: Reserva Nacional Nuble, Parque younger trees, eventually exfoliating in irregular
Nacional Conguillio and Parque Nacional Tolhuaca. small papery strips; bark colour reddish brown,
Germination experiments conducted in Chile and weathering blackish brown. New foliage yellow-
the UK have resulted in increased germination rates ish green, turning dark green. Seedlings and young
that could help restocking lost or diminished natural plants with slender leaves 1220mm long and 1.8
populations. 2.3mm wide; apex acute. Leaves on mature trees
IUCN: VU [B2a, b (iiv)] alternate, twisted at a petiolate base, mostly pecti-
nately arranged in two rows and one plane, linear,
Uses 1020mm long, 2.23.1mm wide, slightly curved
sideways above base, widest above the middle, with
In the past the wood was much used to make furni- a distinct grooved midrib adaxially (upperside) and
ture; at present trees are not felled at a commercial a midrib abaxially (underside); margins entire; apex
scale but its wood can become available when for- obtuse or sometimes acute; leaf colour lustrous dark
est patches that contain it are cleared for other land green above, dull light green below. Stomata in two
uses. The succulent tissue surrounding the seeds is bands of numerous lines separated by midrib on
sweet and these are harvested for consumption by abaxial (under) side of leaves. Pollen cones on lateral
local people, as most female trees bear large quan- and axillary spikes towards ends of foliage branches;
tities of fruit each year. There are also potential up to 20 spreading cones per spike, each cone axil-
medicinal properties, e.g. chemicals that actively lary to a small leaf, 710mm long and 2mm wide.
deter fungal infections detected in the bark of this Microsporophylls sub-peltate with acute apex,
tree. Prumnopitys andina is in cultivation in arbo- 0.5mm wide, incurved at anthesis, light green, each
reta, botanic gardens and private tree collections with two yellow pollen sacs. Seed cones solitary
in many countries, mostly based on cuttings taken on 515mm long branchlets with deciduous scale
leaves, axillary to a scale leaf (bract), consisting of Prumnopitys ferruginea (G. Benn. ex D. Don)
an epimatium enclosing a globose, slightly flattened de Laub., Blumea 24 (1): 190. 1978. Podocarpus
seed, growing into a globose, pruinose pseudo-fruit ferrugineus G. Benn. ex D. Don, in Lambert,
1012mm diam. with an apical crest. Descr. Pinus, ed. 8, 2: [189]. Type: Habitat in
Nova Zelandia. Phillip, Bennett. (LINN?, not
Distribution designated).
Ecology Description
Prumnopitys ferruginoides is a tree of montane rain- Trees to 35m tall; trunk to 1.8m d.b.h., with an erect
forest distributed mainly on ultramafic soils derived bole. First order branches ascending or spreading in
from serpentine; in the extreme north of the island large trees forming a rounded crown. Bark smooth
(Mont Pani) it occurs on micaschist. Its altitudinal on younger trees, becoming scaly and exfoliating in
range is approximately from 150m to 1400m a.s.l. irregular small strips; bark colour purplish brown,
This species is scattered in these forests as mostly weathering grey. Seedlings and young plants with
solitary trees dispersed by birds, which eat the slender, nearly straight to more or less falcate leaves
pseudo-fruits, and therefore this species can occur 2030mm long and 24mm wide; apex acute.
randomly with many other trees, both conifers and Leaves on mature trees alternate, twisted at a peti-
angiosperms. Apparently this species is shade tol- olate base, mostly pectinately arranged in two rows
erant, juveniles up to 56m tall growing up under and one plane, linear, (8)1520mm long, 23.5mm
canopy have markedly larger leaves than mature wide, slightly curved sideways above base, mostly
trees of which the crowns have reached abundant parallel-sided, with a distinct grooved midrib adaxi-
sunlight. ally (upperside) and a thin midrib abaxially (under-
side); margins entire; apex mostly obtuse or slightly
Conservation acuminate; leaf colour green above, glaucous below.
Stomata in numerous lines not separated by mid-
IUCN: LC rib on abaxial (under) side of leaves. Pollen cones
on lateral and axillary, 34cm long spikes towards
Uses ends of foliage branches; 1520cones axillary to
2mm long bracts, 810mm long and 2mm wide.
There are no uses recorded of this species. If it is or Microsporophylls sub-peltate with broadly triangu-
was logged it would have been treated as podocarp lar apex, 0.7mm wide, imbricate, light green, each
timber not distinct from other tree forming species, with two yellow pollen sacs. Seed cones solitary on
but its scarcity in the forest means it has no economic 1.52.5cm long axillary branchlets with persistent
scale leaves at base and deciduous reduced leaves, database, which does not include all relevant major
subtended by a longer scale leaf (bract); consisting of herbaria). The Conifer Specialist Group believes it
an epimatium enclosing an ovoid, slightly flattened, is prudent to flag this species as NT as deforestation
rugose seed, growing into a subglobose, pruinose may well push it into a threatened category in future.
pseudo-fruit 10mm long and 8mm wide with a This species has been recorded from the Machu
small apical crest. Pichu National Park in Peru.
IUCN: NT
Taxonomic notes
Uses
Pilger (op. cit.) mentioned collections by Karsten
(unspecified) and A. Fendler (No. 1289) when The timber of Prumnopitys harmsiana is valued for 949
describing Podocarpus harmsianus as a new spe- the construction of houses and large trees are often
cies from Venezuela. When De Laubenfels (op. cit.) selectively logged for that purpose. It is probably
transferred this species to the genus Prumnopitys, he the tallest of the Andean species in the genus and,
did not designate its type; that opportunity is here depending on its forest setting, may grow a sizable
fulfilled with a duplicate of Fendler 1289 at K desig- straight bole. This species is not in cultivation out-
nated as the lectotype. It consists of a foliage branch side a few arboreta and botanic gardens in (sub)
of a probably juvenile (sterile?) treelet. Pilger saw tropical countries.
the spiked pollen cones, apparently on other mate-
rial which was likely destroyed at Berlin-Dahlem (B)
during World War II, and stated not to know the seed Prumnopitys ladei (F. M. Bailey) de Laub., Blumea
cones. In describing Podocarpus utilior from Peru 24 (1): 190. 1978. Podocarpus ladei F. M. Bailey,
two years later, Pilger (op. cit.) had seed cones but Queensland Agric. J. 15 (8): 899, t. 22. 1905. Type:
no pollen cones. His suspicion that the two might be Australia: Queensland, [Mount Spurgeon, Mitchell
one species has been confirmed to be the case and River, Port Douglas, F. W. H. Lade, leaf specimens,
the two descriptions can be united. December 1902; immature fruiting specimens,
May 1905.] (syntypes, BRI). Fig. 299, 300
Distribution
Etymology
The Andean regions of Bolivia, Colombia, Equador,
Peru, and Venezuela. The species epithet commemorates F. W. H. Lade,
TDWG codes: 82 VEN 83 BOL CLM ECU PER who collected the type specimens in 1902 and 1905.
Prumnopitys harmsiana is widespread but relatively Mount Spurgeon Black Kauri Pine
uncommon in the Andean lower montane to mon-
tane forest belt at altitudes between 1000m and Description
2200m a.s.l.
Trees to 33m tall; trunk to 1m d.b.h., may be but-
Conservation tressed; upper part of tree densely branched form-
ing a rounded crown. Bark more or less smooth
This species has a wide distribution in a forest type on younger trees, becoming scaly and breaking up
that is under pressure throughout its range as a into thin flakes on the bole of large trees; inner bark
result of deforestation. Most herbarium collections fibrous; bark colour reddish brown, weathering grey.
are from Bolivia and Peru. The extent of occurrence Leaves on young trees alternate, twisted at a short
(EOO) has not been calculated due to a lack of speci- decurrent and more or less petiolate base, mostly
men locality data in the other countries, where it is pectinate, (nearly) straight, 1525mm long, 34mm
believed to occur more frequently than the avail- wide, with a distinct, raised midrib on both sides;
able data suggest (data largely from the Tropicos margins entire, parallel to shortly below the obtuse
and minutely hyaline apex; leaf colour green to P. M. Hyland 7882); this implies that logging of pri-
brownish red. Leaves on adult trees shorter, mostly mary forest was then still undertaken and it would
pectinate, linear or more or less oval in the shortest not have spared this species. Several of these logging
leaves, (5)812(16)mm long and 2.53.5mm wide, areas existed there between 19601980 according to
with a faint midrib on both sides; margins slightly data on herbarium labels. On the other hand, recent
incurved; apex obtuse or faintly apiculate; leaf colour surveys have extended the occurrence of P. ladei to a
green above, whitish green below. Stomata densely few localities in between both mountains.
scattered on both sides of leaves, not separated by IUCN: VU (D1, D2)
midrib. Pollen cones axillary, solitary, sessile or on
a short peduncle, oblong, 79mm long and 23mm Uses
950 wide. Microsporophylls sub-peltate with obtuse
apex, 1mm wide, light green, each with two yellow The wood of this species is used for construction and
pollen sacs. Seed cones solitary on short axillary carpentry, but it is too rare to be of much economic
branchlets with deciduous small leaves, terminal importance. It is said to have potential as an orna-
with 12 bracts, the upper bract subtending a single mental tree but is rarely used in that capacity and
ovule (seed). Epimatium enclosing a broadly ovoid, may not be present outside botanical collections.
slightly flattened brown seed, growing into an ellip-
soid to globose, green or blue-green and finally dark
blue to blue- black pseudo-fruit 2230mm long and Prumnopitys montana (Humb. & Bonpl. ex Willd.)
1825mm wide, with a distinct apical protrusion. de Laub., Blumea 24 (1): 189. 1978. Taxus montana
Humb. & Bonpl. ex Willd., Sp. Pl. 4 (2): 857. 1806;
Distribution Podocarpus montanus (Humb. & Bonpl. ex Willd.)
Lodd. ex Britton, Bull. Torrey Bot. Club 16: 13.
Australia: NE Queensland, (Mt. Lewis, Mt. Spurgeon 1889. Type: [Habitat in montibus Peruvianis.]
and a few other localities). (Humboldt & Bonpland in P-Bonpl.?, n.v.).
TDWG codes: 50 QLD-QU
Podocarpus montanus (Willd.) Lodd. var. diversifo-
Ecology lius Dallim. & A. B. Jacks., Handb. Conif.: 51. 1923.
Podocarpus montanus (Willd.) Lodd. var. densifolius
This rare species is restricted to (remnants of) J. T. Buchholz & N. E. Gray, J. Arnold Arbor. 29: 70.
rainforest habitat mainly on two mountains in NE 1948.
Queensland. These mountains are granitic outcrops Podocarpus montanus (Willd.) Lodd. var. meridensis
in the Atherton Tableland. The altitudinal range of J. T. Buchholz & N. E. Gray, J. Arnold Arbor. 29: 71.
the species is between 930m and 1400m a.s.l. The 1948.
rainforest on these mountains is dominated by
angiosperms with scattered podocarps: Podocarpus Etymology
smithii, Prumnopitys ladei and Sundacarpus amarus.
Prumnopitys ladei is the rarer of these species. The The species epithet refers to its habitat in mountains.
seeds are eaten by native rats.
Vernacular names
Conservation
diablo fuerte, pino criollo, pino de montaa, romer-
The surviving rainforests on both mountains are illo hembra (Spanish)
now partly within protected areas. Logging has
almost certainly reduced the area of occupancy Description
(AOO) of Prumnopitys ladei on Mt. Lewis, pres-
ently the smaller of the two populations. In 1974 a Trees to 30m tall; trunk to 1.5m d.b.h., with an erect
herbarium collection was made within the North and straight bole, commonly buttressed at base. First
Mary Logging Area of State Forest Reserve 143 (B. order branches spreading or ascending in large trees
forming a broadly domed or rounded crown. Bark line Sphagnum bogs prevail, with the scrub vegeta-
smooth on younger trees, becoming scaly and exfo- tion retreating to drier spots.
liating in irregular small strips; bark colour purplish
brown, weathering grey. New foliage yellowish or Conservation
glaucous green, turning dark green. Seedlings and
young plants with slender leaves 1220mm long and Logging has caused a reduction of population size of
1.82.3mm wide; apex acute. Leaves on mature trees at least 30% and the decline continues.
alternate, twisted at a petiolate base, mostly pecti- IUCN: VU (A2a, c, d)
nately arranged in two rows and one plane, linear,
(8)1220mm long, 23.5mm wide, slightly curved Uses
sideways above base, widest above the middle, with 951
a distinct grooved midrib adaxially (upperside) and This podocarp is a very valuable timber tree; the
a midrib abaxially (underside); margins entire; apex wood is of medium density, straight grained, work-
acuminate or sometimes obtuse; leaf colour lustrous able and durable. It is used for construction, general
green above, glaucous below. Stomata in numerous carpentry, indoor floors, cabinet making, and tool
lines not separated by midrib on abaxial (under) handles. It is also very good for wood turning with
side of leaves. Pollen cones on leafless or nearly leaf- light yellow sapwood and reddish heartwood. Where
less, lateral and axillary spikes towards ends of foli- the forest has been cleared to make way for pasture,
age branches; up to 25 spreading cones per spike, individual trees or small groups are often left stand-
510mm long and 2mm wide. Microsporophylls ing to provide shade for cattle. It is rare in cultiva-
sub-peltate with acute apex, 0.5mm wide, spreading tion, mainly restricted to some botanic gardens.
at anthesis, light green, each with two yellow pollen
sacs. Seed cones one or two on 23cm long branch-
lets with leaves, axillary to a scale leaf (bract); con- Prumnopitys standleyi (J. T. Buchholz & N.
sisting of an epimatium enclosing an ovoid, slightly E. Gray) de Laub., Blumea 24 (1): 190. 1978.
flattened, rugose seed, growing into an ovoid, olive- Podocarpus standleyi J. T. Buchholz & N. E. Gray,
green ripening to yellow pseudo-fruit 1416mm J. Arnold Arbor. 29: 72. 1948. Type: Costa Rica:
long and 1012mm wide with an apical knob. Alajuela, Poas Mts., Volcan Poas, A. Tonduz 10333
(holotype US).
Distribution
Etymology
The Andean regions of Colombia, Equador, Peru
and Venezuela. This species was named after the botanist Paul C.
TDWG codes: 82 VEN 83 CLM ECU PER Standley (18841963), who studied the flora of Costa
Rica.
Ecology
Vernacular names
Prumnopitys montana is a species of high mon-
tane forest and timberline scrubland, ranging from Ciprecillo (Spanish).
1500 to 3600m a.s.l. In the tropical high montane
rainforest (cloud forest) it is a constituent in a for- Description
est type dominated by angiosperms such as Clusia
(Guttiferae), Weinmannia (Cunoniaceae), Ocotea, Trees to 25(30) m; trunk monopodial, to 1.5(2)
and various other genera in Lauraceae. At the high- m d.b.h. Bark smooth in young trees, becoming
est altitudes this forest is replaced by scrub, often scaly on large trunks, exfoliating in flakes, brown
dominated by Ericaceae and Podocarpaceae, where weathering grey. Branches spreading, forming a
P. montana is an emergent but low tree or a dwarfed rounded crown, foliage branches alternate, form-
tree. Both vegetation formations are rich in epi- ing flat sprays. Foliage twigs slender, glabrous, ini-
phytes. In places with high water table near the tree tially covered in narrow decurrent leaf bases, later
with thin, exfoliating bark. Leaves alternate, usually with 0.11 per ha according to one study. The tree
pectinately arranged in two rows and one plane, is selectively logged, while only a few localities are
linear, twisted and slightly curved above the decur- within protected areas. Some collections were made
rent base and sometimes near apex, or more or less within the following protected areas: Volcan Poas
falcate, (12)1520(25)mm long, 23.5mm wide, Reserve and Rio Macho Reserve.
narrowed towards the petiolate base; apex acute to IUCN: EN [B1a, b (iiii, v)]
mucronate. Adaxial leaf surface (upperside) with a
conspicuous longitudinal groove through the mid- Uses
dle, dark green; abaxial surface (underside) with a
narrow longitudinal midrib, with very small stomata The timber of Prumnopitys standleyi is valued for
952 in numerous lines covered with a greyish white wax. the construction of houses and large trees are often
Pollen cones numerous on slender, leafless, axillary selectively logged for that purpose. The wood is suit-
spikes towards ends of foliage sprays, close together, able for construction, flooring, veneer, furniture
each cone on a 23mm long stalk (bare base of and also used as fenceposts and for firewood. This
the rachis), cylindrical, 1015mm long, 22.5mm species is not in cultivation outside perhaps a few
wide at anthesis. Microsporophylls numerous, heli- botanic gardens.
cally arranged, sub-peltate, curved upwards with a
scarious-denticulate, acute apex, each bearing near
base two relatively large pollen sacs. Seed cones Prumnopitys taxifolia (Banks & Sol. ex D. Don)
terminal on small 0.51.5cm long branchlets with de Laub., Blumea 24 (1): 190. 1978. Dacrydium
a few reduced leaves; producing a single subapical, taxifolium Banks & Sol. ex D. Don, in Lambert,
broadly oval pseudo-fruit ca. 10mm long, 78mm Descr. Pinus, ed. 1, 2: 25. 1824. Type: New Zealand:
wide, consisting of a fleshy epimatium enclosing a [locality not stated], J. Banks & D. Solander s.n.
broadly ovoid, slightly flattened seed, initially green, (holotype BM.
bluish when mature.
Etymology
Distribution
The species epithet refers to the yew-like leaves (yew
Costa Rica (Alajuela, Cartago, Heredia, San Jos). = Taxus).
TDWG codes: 80 COS
Vernacular names
Ecology
Black pine; matai (Maori)
Prumnopitys standleyi is a climax forest species in
montane tropical rain forest. Its altitudinal range Description
(GIS data) is 1349m to 2603m a.s.l.; based on her-
barium specimen data it is 18002960m, while some Trees to 25m tall; trunk to 1.3m d.b.h., crown wide,
reports mention altitudes above 3000m. Annual domed or rounded. First order branches ascending
rainfall is high, from 2000mm to 4000mm. The and eventually spreading in old trees opening up
species is reported to be slow-growing. the wide spreading crown. Bark smooth on younger
trees, becoming scaly and exfoliating in irregular
Conservation flakes leaving multi-coloured patterns on the bole.
Flakes on younger trees large, becoming much
This species is restricted to four to five locations. smaller on large boles of older trees, when longitu-
The extent of occurrence (EOO) based on specimen dinal grooves appear; bark colour pattern varying
distribution data is calculated to be 611 km2 and the from greenish grey or nearly white to reddish or
area of occupancy (AOO) 130 km2. Regeneration is purplish brown. New foliage bright green, turning
reported to be poor and trees are scarce in the forest, dark green. Seedlings and young plants with long,
whip-like shoots and brownish leaves 510mm Distribution
long and 12mm wide. Leaves on mature trees
alternate, twisted at a petiolate base, mostly spread- New Zealand: North Island, South Island.
ing to all sides but on some shaded branchlets more TDWG codes: 51 NZN NZS
pectinate, linear, (6)1020mm long, (1)1.52mm
wide, slightly curved backward or assurgent, or Ecology
nearly straight, with a distinct, slightly raised mid-
rib on both sides; margins entire; apex acuminate or Prumnopitys taxifolia is a constituent species of low-
sometimes obtuse; leaf colour lustrous green above, land to montane (20800m or even 1000m a.s.l.)
whitish green below. Stomata in two broad bands podocarp (conifer) forest, dominated by Podocarpus
separated by a green midrib on abaxial (under) side totara, Dacrycarpus dacrydioides, or Dacrydium 953
of leaves. Pollen cones on leafless or nearly leafless, cupressinum, and with Prumnopitys spp., Manoao
lateral and axillary spikes towards the ends of foli- colensoi, Halocarpus kirkii, and Phyllocladus tricho-
age branches; up to 25 spreading cones per spike, manoides as frequent associated conifers. In low-
512mm long and 2mm wide. Microsporophylls land areas few undisturbed forest remains and here
sub-peltate with rounded apex, 0.5mm wide, light numerous angiosperms are often mixed in, while at
green, each with two bright yellow pollen sacs, giv- higher altitude especially in South Islands Westland
ing the cones a yellow colour when mature. Seed District the genus Nothofagus is often codominant.
cones several on stout branchlets with deciduous As a slow growing conifer, reaching a maximum
scale leaves, axillary to a scale leaf (bract) and con- age in excess of 1000 years (Enright & Hill, 1995)
sisting of an epimatium enclosing an ovoid, slightly Prumnopitys taxifolia is a species that typically regen-
flattened, rugose seed, growing into a globose erates only after episodal distrurbances such as fires
or nearly spherical, purplish black pseudo-fruit caused by volcanic eruptions. It then competes with
810mm diam. after most of the undeveloped, other invading podocarps as well as angiosperms for
green seeds have fallen. light and space, resulting in the long term survival
of only a few individuals of this species in the forest.
Taxonomic notes
Conservation
The occurrence of pollen cones in spikes, i.e. situated
on what appear to be specialized axillary shoots with- IUCN: LC
out functional leaves and arranged in rows, is rare in
conifers. It occurs in Metasequoia, Taxodium (both Uses
Cupressaceae) and in five species of Prumnopitys:
P. andina, P. exigua, P. montana, P. standleyi, and P. Matai is a slow growing tree and produces dense, hard
taxifolia (Podocarpaceae). It is recorded from the and heavy brown wood with exceptional strength
fossil record in Cordaitales, an extinct order of gym- and durability. It was extensively logged and used for
nosperms that was perhaps ancestral to conifers, railway sleepers, bridges, house building, especially
and its limited occurrence in two not closely related floors and weatherboards, and to a lesser extent for
extant conifer families indicates that this character furniture. The logging of this and other native coni-
has evolved more than once. There may be some fers is now outlawed and consequently the use of its
adaptive advantage, because these elongated shoots wood has greatly diminished. It is rare in cultivation
bearing pollen cones without leaves help to expose and only suitable for regions with mild winters that
the former unimpeded to the air, so that the slightest rarely experience light frosts and have plentiful and
breeze could disperse the pollen. evenly distributed rainfall.
Pseudolarix Gordon, Pinetum: 292. 1858 (nom. cons.). Type: Pseudolarix amabilis
(J. Nelson) Rehd. [Larix amabilis J. Nelson] (Pinaceae).
Laricopsis Kent, in Veitch, Man. Conif., ed. Kent: Pseudolarix amabilis (J. Nelson) Rehd., J. Arnold
403. 1900 (nom. illeg.). Type: Laricopsis kaempferi Arbor. 1: 53. 1919. Larix amabilis J. Nelson,
(Lindl.) Kent [Abies kaempferi Lindl.]. Chrysolarix Pinaceae: 84. 1866; Chrysolarix amabilis (J. Nelson)
H. E. Moore, Baileya 13: 133. 1965. Type: Chrysolarix H. E. Moore, Baileya 13: 133. 1965. Type: United
amabilis (J. Nelson) H. E. Moore [Larix amabilis Kingdom: England, cultivated, G. Gordon s.n.
J. Nelson]. (holotype K). Fig. 301, 302
954
Greek: pseudo = false; i.e. resembling but not equal- Abies kaempferi Lindl., Penny Cyclop. 1: 34. 1833,
ling the genus Larix. (quoad descr., non basion.); Pseudolarix kaempferi
(Lindl.) Gordon, Pinetum: 292. 1858.
Description Pseudolarix pourtetii Ferr, Trav. Lab. Forest.
Toulouse T. 1 (4, 4): 1, f. 113. 1944.
See the species description.
Etymology
Taxonomic notes
The species epithet (Latin amabilis = lovely or lov-
The genus Pseudolarix is known from the fossil record able) refers to its golden yellow autumn foliage.
(Early Cretaceous to Pleistocene) in numerous
localities in North America and Eurasia. It occurred Vernacular names
in Europe at least until the end of the Pliocene and
in the Early Tertiary extended far north into the Chinese golden larch; jin qian song (Chinese)
Arctic. At least two species were present during
the Tertiary. It is a classic example of a relict coni- Description
fer which once was very common and widespread,
but has been reduced to a few localities in China. Monoecious, deciduous trees to 3040(68) m
This conifer was traditionally classified with Cedrus tall, d.b.h. to 1.5m; trunk monopodial, straight or
and Larix based on vegetative characters but more curved; crown irregular, broadly domed and open
detailed phylogenetic analyses using both morphol- in old trees. Bark rough and scaly, breaking into
ogy and DNA have refuted this. Its affinities are with thick, square scales, brown-grey. Branches spreading
Keeteleria, Nothotsuga and Tsuga in an abietoid horizontally or slightly ascending, drooping at ends.
clade (Farjon, 1990; Wang et al., 2000) separate from Branchlets slender, flexible, in second year thicker
a pinoid clade which includes among others Cedrus and firm, pale yellowish, darkening to pinkish or
and Larix. The short shoots and deciduous leaves are purplish brown, later grey, ridged and grooved, gla-
parallelisms which evolved at least twice within the brous, young shoots slightly pruinose. Short shoots
Pinaceae. cylindrical-oblong, marked with annual rings of
perular scale bases, length increasing with age, 0.5
Distribution 3.5cm. Vegetative buds of long shoots ovoid, pale
brown, (axillary) lateral buds of short shoots globu-
As for the species. lar, yellowish green, 23mm, not resinous. Leaves
spirally arranged, remote on long shoots, dense
on short shoots, in false terminal whorls of 1030,
assurgent, (2)36(7.5)cm long, (1.5)23(4)mm
wide, linear, flaccid, flattened, shallowly grooved
above, keeled below, obtuse or acutish at apex;
hypostomatic, stomata in 2 broad bands separated
by a midrib; leaf colour light green, pale green below, tree species (Wang, 1961). Its altitudinal range is
turning bright yellow, finally orange-brown in from 180m to 1000m a.s.l., so it is a component of
autumn. Pollen cones terminal on short shoots, clus- lowland forest in a warm temperate, humid climate
tered with 1025 in umbels, ovoid at first, cylindrical which can experience occasional cold winter frost at
and pendulous when ripe, becoming yellow. Seed the higher altitude. This species grows on a variety
cones terminal on short shoots, solitary, erect, with of soils derived from acidic rock; it does not occur
short-leaved, 0.51.5cm long peduncles, ovoid-glo- on limestone. Several localities where P. amabilis still
bose, open, resembling artichokes, (4)57(8)cm occurs in the wild are likely to have been altered by
long, 45.5cm wide with opened scales, colour light man and will support secondary forest, indicated by
green or purplish green, ripening to light yellowish the presence of Cunninghamia lanceolata and Pinus
brown, disintegrating; cone rachis weak, narrowly spp.; some of these trees may have been planted in 955
conical. Seed scales narrowly deltoid-triangular, reforestation projects nearby. On the other hand, P.
auriculate-pedicellate at base, slightly recurved or amabilis is a light demanding tree and perhaps not a
straight at apex, 2.83.5 1.31.8cm at mid-cone, constituent of a late phase in the succession, which
coriaceous, striated or rugose at maturity, sometimes tends to angiosperm (broad-leaved) dominance.
with orange-yellow resin, nearly glabrous; margins Undisturbed primary forest that contains P. amabilis
entire, erose in old scales. Bracts lanceolate, 68mm is now extremely rare and, if it still exists, should be
long, entirely included. Seeds ovoid-cuneate, partly subjected to detailed ecological study.
enveloped by wing membrane, 57mm long, pale
brown or rose-brown; seed wings obliquely pointed, Conservation
extending beyond the seed scales, 2330 813mm,
yellow or pale brown. The natural distribution of this species is based upon
the populations in and around Zhejiang and two
Distribution localities in southern Anhui and northern Jiangxi
and (tentatively) one in Hunan; all other herbarium
China: lower Chang Jiang (Yangtse) valley (Anhui, based localities are believed to be planted or intro-
Hunan, Jiangxi, Jiangsu, Zhejiang). The true geo- duced and here and there perhaps naturalized; they
graphic range of this species is difficult to ascertain. are excluded from the assessment. This species is
In Atlas of the Gymnosperms of China (Ying et al., very rare in the wild and occurs in a few remnants
2004), a map with dots (based on herbarium speci- of primary forest on isolated mountains. Most loca-
men data from Chinese herbaria) is produced which tions are not within protected areas and loss of habi-
gives a much wider range than here presented. A tat is still continuing in this densely populated part
map given in China Plant Red Data Book 1 (Fu & Jin, of China. A detailed survey of existing truly natural
1992) is similarly optimistic. A check by Dr Xiang populations occurring in primary mixed forest to
Qiao-ping in the Beijing Herbarium (PE) revealed determine more exactly the number of remaining
that most of these specimens were collected from mature trees, as well as the number of populations
planted trees or trees in villages and towns (e.g. near and their localities, is a first and necessary step to
temples) that did not occur in the wild. This problem conservation action. This should next lead to the
is acknowledged in Flora of China 4 (1999). Wang establishment of protected areas including the major
(1961) has given an indication where we may expect sites and populations.
this attractive tree to occur in its natural habitat: his IUCN: VU [B2a, b (iii, v)]
examples are all in northern Zhejiang.
TDWG codes: 36 CHS-AH CHS-HN CHS-JS CHS-JX Uses
CHS-ZJ
This species has been planted in arboreta and parks
Ecology in many countries, providing ample material for
study. Herbarium specimens from (wild) locations
Pseudolarix amabilis is a component of the mixed in China are very rare, therefore the majority of
mesophytic, partly evergreen forest, very rich in material studied has come from European arboreta.
Due to its rarity and slow growth the value as a tim- probably centuries before it came to Europe. It is
ber tree of this conifer is limited to local use, pri- now present in most collections of planted coni-
marily for boat building, wooden foot bridges, and fers in the temperate climate regions of the world,
furniture. The Chinese golden larch has been in especially in the northern hemisphere, but it
cultivation in Europe since the early 1860s when it remains relatively uncommon in the trade, perhaps
was first described as a species of larch in England, because it is difficult to propagate and slow grow-
based on small plants grown from seed and from ing. It is mostly used as a specimen tree in large
seedlings transported by Robert Fortune from gardens and parks in W Europe and the eastern
China to England in a Wardian case. It is a desir- USA. A few dwarfed forms have been selected and
able and successful tree in cultivation, which has are grown as cultivars in Chinese and Japanese style
956 been responsible for its wide distribution in China gardens.
Pseudotaxus W. C. Cheng, Notes Forest. Inst. Nat. Centr. Univ. Nanking,
Dendrol. Ser., 1: 1. 1947. Type: Pseudotaxus chienii (W. C. Cheng) W. C. Cheng
[Taxus chienii W. C. Cheng] (Taxaceae).
Nothotaxus Florin, Acta Horti Berg. 14 (9): 394. 1948 whorled, spreading or ascending, forming a bushy
(nom. illeg.). Type: Nothotaxus chienii (W. C. Cheng) crown. Foliage branchlets subwhorled or suboppo-
Florin [Taxus chienii W. C. Cheng]. site, slender, terete, smooth, green turning dark green
in the second year, terminating in small, ovoid-con-
Greek: pseudo- = false, i.e. resembling but not equal- ical buds with persistent bud scales. Leaves helically
ling; Taxus is the classical Latin name for yews. arranged, distichous, spreading at 4090 to shoot 957
axis, well spaced but some overlapping at various
Description angles, 1.22.8cm long, linear, straight or slightly fal-
cate in some leaves only, (1.5)24(4.5)mm wide,
See the species description. short petiolate or sessile, obtusely rounded at base;
margins parallel and revolute, abruptly converg-
Distribution ing and terminating in a mucronate apex; mucro
hooked, pale whitish green; leaf texture slightly cori-
As for the species. aceous, smooth or slightly rugose; leaf colour green
above, abruptly turning dark green in second year,
green with two white bands below. Midrib raised on
Pseudotaxus chienii (W. C. Cheng) W. C. Cheng, both sides, ca. 0.4mm wide, continuous. Stomata in
Notes Forest. Inst. Nat. Centr. Univ. Nanking, two bands 0.51mm wide on abaxial (lower) side,
Dendrol. Ser., 1: 1. 1947. Taxus chienii W. C. Cheng, slightly wider than midrib and green margins, form-
Contr. Biol. Lab. Sci. Soc. China, Sect. Bot. 9: 240. ing 1020 intermittent white lines in each band.
1934; Nothotaxus chienii (W. C. Cheng) Florin, Pollen cones usually concentrated towards ends of
Acta Horti Berg. 14 (9): 394. 1948. Type: China: lateral branchlets, axillary, solitary, sessile, globose,
Zhejiang, Longquan Xian, Maoshan, S. Chen 1384 23.5mm long, 3mm wide, with 4 pairs of decus-
(lectotype PE). Fig. 303 sate basal bracts; microsporophylls 612, decussate,
peltate, with 46 radially placed pollen sacs. Seed-
Pseudotaxus liana Silba, Phytologia 81 (4): 327. bearing structures axillary, solitary, sessile, with 7
1996 [liiana]; Pseudotaxus chienii (W. C. Cheng) pairs of decussate basal bracts. Seed arils cupular,
W. C. Cheng subsp. liana (Silba) Silba, J. Int. Conifer only partly enclosing the seed, succulent and white
Preserv. Soc. 14 (1): 19. 2007. when ripe, 57mm long. Seeds ovoid, 58mm long,
45mm wide, slightly flattened distally, with two
Etymology obscure ridges below mucronate apex.
Abietia A. H. Kent, in Veitch, Man. Conif., ed. Kent: and the North American Cordillera, southward
474. 1900. Type: Abietia douglasii (Sabine ex D. Don) increasingly scattered. Asia: in the eastern Himalaya,
A. H. Kent [Pinus douglasii Sabine ex D. Don]. Central and SE China, N Viet Nam, Taiwan,
Japan.
Greek: pseudo = false, i.e. resembling but not equal-
ling the genus Tsuga. Key to the species and some varieties of
Pseudotsuga 959
Description
Monoecious, sometimes very tall (100m), ever- 1a. Seed scales not wider than long, thin; bracts
green trees, monopodial with a columnar trunk and much longer than the seed scales, usually not
a conical or spreading crown and pseudo-whorled, reflexed. Leaves 11.5mm wide, parted above
plagiotropic branching (Massarts model). Resin the shoot but not pectinate. Bark on the lower
canals in wood, bark, leaves and seed cones. Bark trunk of large trees very thick and deeply fis-
thick, with deep, longitudinal fissures in large trees. sured P. menziesii
(Sub-)terminal buds ovoid conical, not resinous, 1b. Seed scales wider than long, relatively thick;
with triangular scales persisting 12 years. Leaves bracts as long as or slightly longer than the seed
spirally arranged, spreading more or less radially scales, often reflexed. Leaves 1.52 mm wide,
or pectinate, placed on small depressions, linear, mostly pectinate. Bark on the lower trunk rela-
flattened, with two bands of stomata only on the tively thin 2
abaxial side. Pollen cones axillary, solitary, catkin- 2a. Seed cones ovoid-cylindrical, very large
like, 12cm long, subtended by conspicuous perular (918cm long); bracts slightly longer than the
scales; microsporophylls spirally arranged, peltate, seed scales, usually not reflexed. Leaf apex
with two pollen sacs containing globular pollen with acute P. macrocarpa
3converging ridges on a smooth surface. Seed cones 2b. Seed cones ovoid, smaller than 8 cm; bracts
axillary, solitary, more or less erect at pollination but equally long as the seed scales, reflexed. Leaf
soon pendulous on curved peduncles, deciduous apex usually emarginate 3
some time after seed dispersal or semi-persistent. 3a. Seed cones broad, ovoid, usually 4.57.5
Bract scales exserted, as long as or longer than the 45.5cm. Leaves variable in length, up to 5cm
seed scales, trilobate, with the cusp longer than the long (leaves 0.72cm long and 23mm wide =
lateral lobes, straight or reflexed. Seed scales with a P. sinensis var. brevifolia)
rounded upper margin and a broad base, opening P. sinensis var. sinensis
more or less wide. Seeds held in a shallow membra- 3b. Seed cones smaller (not longer than 5 cm).
nous cup covering one side of the seed and which is Leaves up to 3cm long 4
continued in an adnate, short, obliquely ovate wing. 4a. Seed cones ovoid-conical; seed scales variable,
Seedlings with (5)710(14) cotyledons. slightly wider than long, with rounded upper
margin P. japonica
4 species. 4b. Seed cones ovoid-oblong; seed scales reniform
to broad flabellate, much wider than long, with
Distribution truncate upper margin
P. sinensis var. gaussenii
North America: from SE Alaska (coast) and British
Columbia to Pueblo, Mexico in coastal mountains
Pseudotsuga japonica (Shiras.) Beissn., Mitt. entire, incurved; base shortly narrowed. Bracts
Deutsch. Dendrol. Ges. 1896 (5): 62. 1896. oblong-spathulate, with 3-lobed apex, central cusp
Tsuga japonica Shiras., Bot. Mag. (Tokyo) 9: 84, longer and narrower than lateral lobes, 1.52cm
t. 3. 1895. Type: Illustration in Bot. Mag. (Tokyo) long, widest (8mm) near apex, exserted, recurved
9, t. 3 opp. p. 41. 1895. (lectotype designated here). or reflexed. Seeds ovoid to more or less triangular,
Fig. 304, 305 4 7mm, pale brown, with dark spots; wings ovate,
410mm long, light or dark brown.
Etymology
Distribution
The species epithet refers to Japan, its country of
960 origin. Japan: W Honshu (Chugoku District), Shikoku.
[Wilsons reporting it from Kyushu has not been
Vernacular names confirmed.]
TDWG codes: 38 JAP-HN JAP-SH
Japanese Douglas-fir; toga-suwara, goyo-toga
(Japanese) Ecology
Etymology Distribution
The species epithet means with large fruit and refers USA: S California (San Rafael Mts., San Gabriel
to the seed cones. Mts., San Bernardino Mts., San Jacinto Mts., and
Santa Ana Mts.).
Vernacular names TDWG codes: 76 CAL
Chinese Douglas-fir; huang shan (Chinese) China: S Anhui, Fujian, SW Guangxi, Guizhou,
W Hubei, N Hunan, N Jiangxi, S Shaanxi, Sichuan,
Description SE Xizang [Tibet], Yunnan, Zhejiang; Taiwan;
N Viet Nam.
Trees to 50m tall, d.b.h. to 2m; trunk more or less TDWG codes: 36 CHC-GZ CHC-HU CHC-SC CHC-
straight, columnar, or forked. Bark on trunk rough YN CHN-SA CHS-AH CHS-FJ CHS-GX CHS-HN CHS-
and very scaly, longitudinally fissured below, grey. JX CHS-ZJ CHT 38 TAI 41 VIE
Branches spreading wide, nearly erect near the top,
forming a domed or flat topped crown. Branchlets Ecology
slender, reddish brown in the first year, soon becom-
ing grey, variably but usually minutely pubescent Pseudotsuga sinensis is a species occurring in low to
in the grooves, soon glabrous, with small, slightly medium high mountains at various elevations. In
elevated, circular or angular leaf scars. Vegetative SE China it occurs between 600m and 1200m, in
buds ovoid or ovoid conical, 48 34.5mm, not or Taiwan between 1000 and 2700m, in Sichuan and
only slightly resinous; bud scales triangular, acute, Yunnan it may be found above 3000m, the high-
lustrous red-brown, deciduous in the second year. est record is 3300m a.s.l. The soils in the SE are red
Leaves more or less remote, pectinate, (0.7)2.5 and yellow earth, in the western part of the range
described as mountain red earth (Wang, 1961). It Description
requires a moist, temperate or warm temperate
climate with annual precipitation between 1000 and Leaves 1.34cm long, 1.52cm wide. Buds ovoid-
2000mm. It is a constituent of the mixed meso- conical, more or less acute, 47 34mm. Seed
phytic forest formation in SE China, mainly with cones 4.58cm long, 35cm wide when opened.
broad-leaved trees, in Sichuan also of the evergreen Seed scales suborbicular to rhombic-orbicular.
oak and deciduous hardwood forest, where besides
Pseudotsuga other conifers occur (e.g. Tsuga chinen- Distribution
sis, T. dumosa, Picea brachytyla var. complanata).
Unlike in North America, Pseudotsuga in Asia does China: S Anhui, Chongqing, Fujian, Guizhou, W
not form extensive stands or occur in pure or nearly Hubei, N Hunan, N Jiangxi, S Shaanxi, Sichuan, 965
pure conifer forests. Its habit as a mature forest tree Yunnan, Zhejiang; Taiwan; N Viet Nam.
reflects this: trees are not columnar or pyramidal but TDWG codes: 36 CHC-CQ CHC-GZ CHC-HU
develop spreading crowns much like the surround- CHC-SC CHC-YN CHN-SA CHS-AH CHS-FJ CHS-HN
ing broad-leaved trees. CHS-JX CHS-ZJ 38 TAI 41 VIE
Uses Conservation
Chinese Douglas fir is a timber tree used for con- A continuing decline, especially of large mature trees
struction, bridge building, furniture and wood fiber. due to logging, has led to a situation in which large
Very large trees are rare as they have been exten- trees have become scarce and most are restricted to
sively logged in the past; hence the economic value inaccessible mountain ridges and summits. Even
of this species has diminished as it appears not very under the broader taxonomic circumscription here
suitable for plantation forestry. Although intro- employed, it has become a rare, though still wide-
duced in Europe under its various synonyms by E. spread tree. Only small subsets of the total popula-
H. Wilson, George Forrest, Camillo Schneider, and tion are within protected areas.
others in the early 20th century, the species remains IUCN: VU (A2c, d)
very rare in cultivation as an amenity tree and is
virtually restricted to a few arboreta and botanic
gardens. Pseudotsuga sinensis Dode var. brevifolia (W. C.
Cheng & L. K. Fu) Farjon & Silba, Phytologia 68:
3 varieties are recognized: 71. 1990. Pseudotsuga brevifolia W. C. Cheng & L.
K. Fu, Acta Phytotax. Sin. 13 (4): 83. 1975. Type:
China: Guangxi, Nanning, near Longzhou, Chinese
Pseudotsuga sinensis Dode var. sinensis. Type: collector No. 40103 (holotype PE).
China: Yunnan, Dali Baizu Zizhizhou, Dengchuan,
E. E. Maire s.n., 1911 (holotype P). Fig. 309, 310 Description
Pseudotsuga wilsoniana Hayata, Icon. Pl. Formos. 5: Leaves 0.71.5(2)cm long, 23(3.2)mm wide.
204, t. 15. 1915; Pseudotsuga sinensis Dode var. wilsoni- Buds ovoid, obtuse, 45 3mm. Seed cones 3.7
ana (Hayata) L. K. Fu & Nan Li, Novon 7 (3): 263. 1997. 6.5cm long, 34cm wide when opened. Seed scales
Pseudotsuga forrestii Craib, Notes Roy. Bot. Gard. suborbicular to rhombic-orbicular.
Edinburgh 11: 189. 1919; Pseudotsuga sinensis Dode
var. forrestii (Craib) Silba, Phytologia 68: 72. 1990. Taxonomic notes
Pseudotsuga xichangensis C. T. Kuan & L. J. Zhou, Fl.
Sichuan. 2: 54. 1983. Pseudotsuga sinensis var. brevifolia has been reported
Pseudotsuga shaanxiensis S. Z. Qu & K. Y. Wang, from the limestone region in northern Viet Nam,
Acta Bot. Bor.-Occid. Sin. 8 (2): 129. 1988. but more recent assessments of the Vietnamese
trees (e.g. Nguyen Tien Hiep et al., 2004) have con- true extent and rarity remains unknown (see under
cluded that the common species on the ridge tops Taxonomic notes). Trees are often inaccessible on
and summits of the limestone mountains is P. sinen- mountain summits but those on easier sites have
sis with normally long leaves. I confirmed this on a often already been logged.
visit to Bat Dai Son in 2002 at least for that area. The IUCN: VU (A2c, d)
distribution of P. sinensis var. brevifolia in Atlas of
the Gymnosperms of China (Ying et al., 2004) on
pp. 100101 borders the distribution of P. sinensis in Pseudotsuga sinensis Dode var. gaussenii (Flous)
Viet Nam given on a map in Nguyen Tien Hiep et al. Silba, Phytologia 68: 71. 1990. Pseudotsuga gausseni
(2004). Is var. brevifolia restricted to China? It seems Flous, Bull. Soc. Hist. Nat. Toulouse 69: 417. 1936.
966 unlikely, but it may also be incorrect determination Type: China: Zhejiang, between Ping-yung and
of specimens on either side of the border that has Taiyuan, R. C. Ching 2144 (holotype NY).
lead to this parapatric distribution pattern. A criti-
cal evaluation of the material (and this taxon) seems Description
needed.
Leaves 1.63cm long, 1.72mm wide. Buds ovoid-
Distribution conical, more or less acute, 45 3mm. Seed cones
3.55.5cm long, 23.5cm wide when opened; seed
China: SW Guangxi, Guizhou (Anlong, Lipuo); N scales broad flabellate to reniform.
Viet Nam (?).
TDWG codes: 36 CHC-GZ CHS-GX Distribution
Decussocarpus de Laub., J. Arnold Arbor. 50: 340. 2a. Adult leaves (5)811 mm long, 23.5 mm
1969 (nom. illeg., Art. 52.1). Type: Decussocarpus nagi wide; midrib on both sides of leaves narrow
(Thunb.) de Laub. [Myrica nagi Thunb.]. and conspicuous R. piresii
2b. Adult leaves (6)1025(40) mm long, 2.55
Latin: retro- = backwards, reversed; Greek: phyl- (7)mm wide; midrib on the adaxial (upper)
los = leaf; referring to the peculiar phyllotaxis (see side of leaves inconspicuous or wide and low 967
description). 3
3a. Midrib on both sides of leaves wide and low.
Description Microsporophylls of pollen cones triangular,
acute R. comptonii
Dioecious, evergreen, dwarfed to large trees. Bark 3b. Midrib inconspicuous on the adaxial (upper)
smooth, exfoliating in longitudinal flakes. Resin side of leaves, narrow and conspicuous on the
canals in leaves and seed cones. Leaves spirally abaxial side. Microsporophylls of pollen cones
inserted on shoots, lanceolate to narrowly ovate, triangular, acuminate 4
obliquely directed into subopposite and decussate 4a. Adult leaves (6)1018(25)mm long. Pollen
apparent pairs by at ca. 90 twisted petioles in oppo- cones 57 22.5 mm. Seeds including the
site directions, forming regular pectinate rows on epimatium 1825(30) 1318(20)mm
lateral shoots. Stomata on both faces of the leaves. R. rospigliosii
Pollen cones axillary or sometimes terminally and 4b. Adult leaves (10)1525(40) mm long.
single or in groups on short, naked peduncles; pol- Pollen cones 1025 22.5mm. Seeds includ-
len bisaccate. Seed cones axillary, solitary or some- ing the epimatium 1420 1013mm
times in pairs, at maturity consisting of a few small, R. vitiense
fused basal bracts and a single large seed covered by a
fleshy, drupe-like, elliptic to ovoid-pyriform epima-
tium colouring deep red, violet or purplish brown. Retrophyllum comptonii (J. T. Buchholz) C. N.
Page, Notes Roy. Bot. Gard. Edinburgh 45: 380.
5 species. 1989. Podocarpus comptonii J. T. Buchholz, Bull.
Mus. Hist. Nat. (Paris), sr. 2, 21: 284. 1949;
Distribution Decussocarpus comptonii (J. T. Buchholz) de Laub.,
J. Arnold Arbor. 50: 344. 1969; Nageia comptonii
Malesia: Maluku [Moluccas], New Guinea, New (J. T. Buchholz) de Laub., Blumea 32 (1): 211. 1987.
Britain, New Ireland; SW Pacific: New Caledonia, Type: New Caledonia: Grande Terre, Province Sud,
Vanuatu, Fiji. South America: W Venezuela, Mt. Mou, J. T. Buchholz 1684 (holotype ILL).
Colombia, Equador, Peru, W Brazil. Fig. 311, 312
SW Pacific: New Caledonia (Plaine des Lacs). There are no uses recorded for this species. It is in
TDWG codes: 60 NWC cultivation in a few botanic gardens as a pot-grown
young plant.
Ecology
This is the only true rheophyte among gymno- Retrophyllum piresii (Silba) C. N. Page, Notes Roy.
sperms. It grows along the banks of small rivers and Bot. Gard. Edinburgh 45: 380. 1989. Decussocarpus
shores of small lakes in shallow water over skeletal piresii Silba, Phytologia 54: 461. 1983; Nageia piresii
970 soils derived from serpentine, ironstone and laterite (Silba) de Laub., Blumea 32 (1): 211. 1987. Type:
with high contents of heavy metals. The maximum Brazil: Rondonia, Serra dos Pacahas Novos,
altitude is 240m a.s.l. The buoyant seeds are easily [in the National Park], N. A. Rosa & J. M. Pires 856
transported by running water and germinate in mud (holotype US).
temporarily falling dry with fluctuating water tables
in lakes and streams. The fleshy epimatium is con- Etymology
sumed by birds, and perhaps fish, but also often rots
away, even on the plant. The thick, fibrous bark is This species was named after one of the collectors, J.
perhaps an adaptation against damage from stones M. Pires, who gathered the type collection.
transported in flash floods that may occur after
torrential downpours over the sparsely vegetated Vernacular names
plain discharge into the rivers. There are few other
plants that occupy this habitat; near the Chtes de la None have been recorded.
Madeleine on the Rivire des Lacs Dacrydium guil-
lauminii occurs at the water edge with R. minus, and Description
there are some sedges (Carex sp.).
Trees to 30m tall, to 80cm d.b.h., bole straight
Conservation and erect, clear of branches for 15m or more in
large trees growing in forest. Bark not described.
Retrophyllum minus is a rare species with a limited Branches spreading, forming a rounded crown.
distribution and a very narrowly defined habitat. Foliage branches alternate or subopposite, spread-
In extremely dry periods it is vulnerable to dehy- ing but becoming somewhat pendulous in shaded
dration and small subpopulations could be wiped branches, with a few dispersed, decurrent, spread-
out by brush fires. These fires have increased in fre- ing, small scale leaves on leading shoots and more
quency in recent decades due to increased human numerous and larger, photosynthetically domi-
visits and activity, e.g. pine plantations and geo- nant leaves on lateral branchlets. Leaves on lateral
logical surveys involving road building, on the branchlets spirally inserted but pectinately arranged,
Plaine des Lacs. Much of this plain falls within a appearing opposite, with a decurrent base and a
mining concession. Currently, this species is only 90 twist at the point where the leaf blade abruptly
protected in a single small botanical reserve at the widens and is free from the shoot, twisting in
Chtes de la Madeleine, which covers only one opposite directions on either side of shoot, orien-
small subpopulation. The total number of mature tating blade on one side of shoot with adaxial side
individual plants is unknown, but lies some- uppermost and on other side with adaxial side
where below 2500 and is inferred to be declining downwards. Leaf shape in mature trees (ob)ovate
because of habitat disturbance, destruction and to elliptic, free part (5)811 23.5mm, widest
fragmentation. in or just above the middle, more or less abrubtly
IUCN: EN [B1ab(iii)+2ab(iii); C2a(i)] narrowing to an obtuse or weakly apiculate apex;
midrib narrow and conspicuous on both sides. Conservation
Stomata on both sides (leaves amphistomatic),
numerous but not in regualar lines. Pollen cones Only known from Rondonia (Serra dos Pacahas
not known. Seed cones on lateral branchlets with Novos) and occurring within a national park; new
small leaves, subterminal fertile bract scale-like, 13 localities recently reported have not been verified
2mm, with a single fertilized ovule growing into by examination of herbarium specimens and need
an inverted seed covered by an epimatium. Seeds confirmation.
including the epimatium ellipsoid, with a promi- IUCN: DD
nent thin crest at distal end, 2022 1014mm,
green becoming fleshy and brownish red when ripe;
seed proper 1214 78mm, with a small beak at Retrophyllum rospigliosii (Pilg.) C. N. Page, 971
micropylar end. Notes Roy. Bot. Gard. Edinburgh 45: 380. 1989.
Podocarpus rospigliosii Pilg., Notizbl. Bot. Gart.
Taxonomic notes Berlin-Dahlem 8: 273. 1923; Decussocarpus rospigli-
osii (Pilg.) de Laub., J. Arnold Arbor. 50: 347. 1969;
This species was first described and named as Nageia rospigliosii (Pilg.) de Laub., Blumea 32 (1):
Decussocarpus piresii from a single collection; its 211. 1987. Type: Peru: Pasco, Oxapampa, N. Esposto
taxonomic base is therefore not very convincing. 556 (holotype USM).
The type collection N. A. Rosa & J. M. Pires (et al.
on duplicates in K and MO) 856 (not 586 as pub- Etymology
lished in the protologue!) has duplicates at K, MO
and US (holotype) and appears to differ from those This species was named after Dr C. J. Rospigliosi of
I have seen of R. rospigliosii in its smaller, narrower the Natural History Museum in Lima, Peru.
leaves and its more oblong seeds. Other stated differ-
ences, such as leaf shapes, are more difficult to assess Vernacular names
for lack of more material. On the other hand, this
genus had not been known from Brazil previously pino hayuelo, pino laso, pino de montaa, pino
(the only other South American species, R. rospiglio- romern, romerillo macho (Spanish)
sii (Pilg.) C. N. Page, occurs in Colombia, Ecuador,
Peru, and Venezuela and has a strictly Andean dis- Description
tribution). More material is needed, but as yet lack-
ing, to ascertain the taxonomic status of this species Trees to 45m tall, to 1.8m d.b.h.; bole straight and
more securely. It should at present be provisionally erect, clear of branches for 15m or more in large trees
accepted as distinct. growing in forest. Bark brown, weathering blackish
grey, exfoliating in large flakes. Branches ascend-
Distribution ing or spreading, forming a pyramidal and eventu-
ally rounded crown. Foliage branches alternate or
Brazil: Rondnia (Serra dos Pacahas Novos). subopposite, spreading but becoming pendulous in
TDWG codes: 84 BZN-RO shaded branches, with dispersed, decurrent, spread-
ing, small scale leaves on leading shoots and more
Ecology numerous and larger, photosynthetically domi-
nant leaves on lateral branchlets. Leaves on lateral
No details are known about the habitat of this spe- branchlets spirally inserted but pectinately arranged,
cies other than that its type specimen has been col- appearing opposite, with a decurrent base and a 90
lected at an altitude of 250m. The area appears to twist at the point where the leaf blade abruptly wid-
be well forested viewed via Google Earth satellite ens and is free from the shoot, twisting in opposite
imagery (accessed 13 March 2009). directions on either side of shoot, orientating blade
on one side of shoot with adaxial side uppermost stands have now disappeared or are reduced to
and on other side with adaxial side downwards. Leaf a few trees. It is considered still in decline as log-
shape ovate-lanceolate or ovate-elliptic, longest in ging and forest clearing have left several trees from
juvenile plants, much shorter in top of old trees, also which herbarium collections were taken by David de
shorter at base and end of lateral branchlets, free part Laubenfels in Venezuela standing alone in pasture
(6)1018(25) 35(6)mm, mostly widest just since 1980. This situation is undoubtedly also found
below the middle, tapering to an acute or apiculate in the other countries where this tree occurs and in
apex; midrib narrow and conspicuous on abaxial Peru populations are becoming reduced in number
side, inconspicuous on adaxial side. Stomata on both of mature trees (see e.g. Reynel et al., 2006).
sides (leaves amphistomatic), numerous but not in IUCN: VU (A2a, c)
972 regualar lines. Pollen cones at or near ends of foli-
age branchlets, axillary to leaves or in groups of 35 Uses
together with one terminal, globose when immature,
57mm long, 22.5mm diam. at anthesis; micro- This species is a valuable timber tree and can yield
sporophylls in spirals, triangular, with an acuminate large sizes of sawn timber. Its wood is of very good
apex, each bearing two pollen sacs. Seed cones on quality, straight grained, of medium density, durable
lateral branchlets with small leaves; subterminal fer- and workable. It is much used for construction, car-
tile bract leaf-like, 3 2mm, with a single fertilized pentry, cabinet making, and wood turning. Mature
ovule growing into an inverted seed covered by an trees are often left standing from forest transformed
epimatium. Seeds including the epimatium subglo- into pasture for cattle and serve to give shelter from
bose to ovoid-pyriform, with a short crest at distal sun and heat for the animals. In future they may be
end, 1825(30) 1318(20)mm, green or glaucous planted for the same purpose; to what extent this is
becoming fleshy and dark red when ripe, drying to a already being done is not known.
hard shell; seed proper 1317 912mm, with a nar-
row beak at micropylar end.
Retrophyllum vitiense (Seem.) C. N. Page,
Distribution Notes Roy. Bot. Gard. Edinburgh 45: 380. 1989.
Podocarpus vitiensis Seem., Bonplandia 10: 366.
Colombia, Ecuador, Central Peru, W Venezuela 1862; Nageia vitiensis (Seem.) Kuntze, Revis.
(Merida and Tachira). Gen. Pl. 2: 800. 1891; Decussocarpus vitien-
TDWG codes: 82 VEN 83 CLM ECU PER sis (Seem.) de Laub., J. Arnold Arbor. 50: 342.
1969. Type: Fiji: Western Division, Viti Levu,
Ecology B. C. Seemann 576 (lectotype K, sheet 2/2, here
designated).
Retrophyllum rospigliosii occurs in montane tropical
rainforest, in which it can attain large size. Its alti- Podocarpus filicifolius N. E. Gray, J. Arnold Arbor.
tudinal range is from 1500m to 3300m (3750m in 43: 74. 1962.
Colombia and Peru) a.s.l., so it occurs in wet rainforest
up to high altitude cloud forest or mossy forest. It can Etymology
form more or less extensive pure stands on exposed
sites, but is more often found scattered among angio- The species epithet refers to Viti Levu, the island in
sperms or sometimes with Prumnopitys spp. Fiji from where it was first described.
This timber species is under much pressure from dakua salusalu, kau solo (Viti Levu); mugo (New
logging and most of the formerly quite extensive Guinea, Danau Paniai [Wissel] Lakes)
Description deep red when ripe; seed proper subglobose, 1216
810mm, with a narrow beak at micropylar end.
Trees to 45(60?) m tall, to 1.5(2?) m d.b.h.; large
trees with spur-butresses at base; bole straight and Taxonomic notes
erect, terete, clear of branches for 20m or more in
large trees growing in forest. Bark smooth, eventu- Lectotype collection number designated by De
ally with faint vertical fissures, light to dark brown, Laubenfels in J. Arnold Arbor. 50: 342 (1969). At K
weathering blackish grey, exfoliating in small dis- there are two sheets of Seemann 576, one (sheet 1/2)
integrating longitudinal flakes. Branches ascending is a seedling (complete) and one (sheet 2/2) a branch
or spreading, forming a pyramidal and eventually from a young tree corresponding with the plate in
rounded crown. Foliage branches alternate or sub- the protologue. They cannot be both the lectotype as 973
opposite, spreading but becoming pendulous in they come from two separate plants. The latter sheet
shaded branches, with remotely dispersed, decur- is to be designated as the lectotype.
rent, appressed, small and thin scale leaves on
leading shoots and more numerous and larger, pho- Distribution
tosynthetically dominant leaves on lateral branch-
lets. Leaves on lateral branchlets spirally inserted Malesia: Maluku [Moluccas]; Papuasia: New Britain,
but pectinately arranged, appearing opposite, with New Guinea, New Ireland; Southwest Pacific: Santa
a decurrent base and a 90 twist at the point where Cruz Islands, Fiji Islands.
the leaf blade abruptly widens and is free from the TDWG codes: 42 MOL 43 BIS NWG-IJ NWG-PN 60
shoot, twisting in opposite directions on either side FIJ SCZ
of shoot, orientating blade on one side of shoot
with adaxial side uppermost and on other side with Ecology
adaxial side downwards. Leaf shape ovate-lanceolate
or ovate-elliptic, longest in juvenile plants, much Retrophyllum vitiense is a tall tree of tropical low-
shorter in top of old trees, also shorter at base and land to montane rainforest, usually occurring as
end of lateral branchlets, free part (10)1525(40) scattered individual emergents. In New Guinea it is
35(7)mm, mostly widest below the middle, sometimes frequent in forests dominated by Agathis
especially juvenile forms gradually tapering to an labillardierei and Lithocarpus spp. In the Bismarck
acute, or in short leaves obtuse apex; midrib nar- Archipelago (New Ireland) it was found growing
row and conspicuous on abaxial side, inconspicu- in montane forest with Serianthes sp., Syzygium sp.,
ous on adaxial side. Stomata on both sides (leaves Dacrycarpus imbricatus, Podocarpus sp., and scat-
amphistomatic), numerous but not in regualar tered palms and frequent thickets of bamboo; here
lines. Pollen cones on relatively short subterminal a tree was reported to be 60m tall. Its altitudinal
or lateral branchlets with scale leaves, in groups of range is from near sea level to 1800m a.s.l. In Fiji
23 together with one terminal, ovate when imma- it is a common constituent of lowland to montane
ture, becoming long cylindrical, 1025mm long rainforest containing several other conifers: Agathis
and 22.5mm diam. at anthesis; microsporophylls macrophylla, Dacrycarpus imbricatus, Dacrydium
in ranks of tight spirals, triangular, keeled, with an nausoriense, D. nidulum, and Podocarpus neriifolius,
acuminate apex, each bearing two pollen sacs. Seed as well as various angiosperm trees including palms.
cones on 612mm long lateral branchlets with scale
leaves; subterminal fertile scale(s) slightly longer Conservation
than sterile scale leaves, with a single fertilized ovule
growing into an inverted seed covered by an epima- Despite logging of this and other podocarp trees in
tium. Seeds including the epimatium more or less many areas of its wide range, which undoubtedly
pyriform, with a short crest at distal end, 1420 has led to some reduction of its area of occupancy
1013mm, green or glaucous becoming fleshy and (AOO), since logged forest is often converted to
other land uses that do not bring back these trees,
Retrophyllum vitiense is as yet not considered a
threatened species.
IUCN: LC
Uses
Squamataxus J. Nelson, Pinaceae: 168. 1866 (nom. more irregularly in outer, sun-exposed foliage, lin-
illeg.). Type: Squamataxus albertiana J. Nelson ear or falcate, (10)1525(30) 23.5mm; base
[Saxegothaea conspicua Lindl.]. petiolate, twisted; blade with a thin midrib on both
sides, at upperside yellowish green or more often
Named after Prince Albert of Saxe-Coburg-Gotha lustrous dark green; flushing leaves pinkish red or
(18191861), Prince Consort to Queen Victoria and a yellowish green; apex mucronate. Stomata in two 975
zealous supporter of science and the arts. centrally placed, whitish bands on abaxial (lower)
side, arranged in continuous lines. Pollen cones axil-
Description lary, situated on branchlets below seed cones or sep-
arately, solitary or sometimes in pairs, subtended by
See the species description. 23 bracts, cylindrical, 46mm long, 1.5mm wide
at anthesis; microsporophylls spirally arranged,
Distribution numerous, minute, with two pollen sacs. Seed cones
terminal on short, axillary branchlets with decidu-
As for the species. ous scale leaves, globular with protruding scale tips
at maturity, 912mm diam., light glaucous green,
purplish and fleshy when ripe. Cone scales 1520,
Saxegothaea conspicua Lindl., J. Hort. Soc. London spirally arranged, at first imbricate; proximal part of
6: 258. 1851. Squamataxus albertiana J. Nelson, (mostly) fertile scales swelling on both sides so that
Pinaceae: 168. 1866 (nom. illeg.). Type: Chile: the tips stand out. Seeds 1 per scale (from 2 inverted
[Patagonia], W. Lobb 81 (holotype not located, ovules), enclosed, subglobose, ca. 3mm diam.,
isotype K). Fig. 315, 316 slightly flattened, lustrous tan or yellowish with a
dull hilum.
Etymology
Distribution
The species epithet means remarkable or striking
(English conspicuous from Latin conspicuus). S Argentina: Chubut, Neuqun, Rio Negro; S Chile:
Aisn, Biobio, La Araucania, Los Lagos, Maule.
Vernacular names TDWG codes: 85 AGS-CB AGS-NE AGS-RN CLC-BI
CLC-LA CLC-MA CLS-AI CLS-LL
Prince Albert yew (England); manio, maniu, manio
macho, manio hembra (Chile) Ecology
Greek: skias = canopy, umbel; pitys = pine or fir green, soon turning light brown, grooved longitu-
tree; referring to the verticillate, spreading cladodes, dinally, dimorphic, with long sections alternating
hence umbrella pine. with much shortened, thicker sections, growing
from terminal, conical, obtuse buds 35 24mm
Description with imbricate, thin chartaceous brown scales.
Leaves much reduced, scale- or cataphyll-like, 35
See the species description. 24mm, the distal, free portion scarious, yellow- 977
ish green, soon brown, caducous. Cladodes (phyl-
Distribution loid shoots) mostly on shortened sections of shoots,
axillary to short, triangular scale leaves, arranged
As for the species. helically in pseudowhorls of (8)1025(40),
spreading, linear, mostly straight, (3)810(13)cm
long, 23.5mm wide, with a narrow longitudinal
Sciadopitys verticillata (Thunb.) Siebold & Zucc., groove from base to apex adaxially and a wider,
Fl. Japon. 2 (1): 3, t. 101102. 1842. Taxus verticil- deeper groove from base to apex abaxially; mar-
lata Thunb., in Murray, Linn. Syst. Veg., ed. 14: 895. gins thick, entire, distally converging to retuse or
Mai-Jun 1784. Type: Japan: Honshu, [locality not emarginate apex; surface smooth, lustrous light to
stated], C. P. Thunberg UPS 23787 (holotype UPS). dark green; abaxial groove lined with white papillae
Fig. 317, 318 concealing the stomata, which are confined to this
groove. Pollen cones in terminal or sublateral clus-
Etymology ters of 820, helically arranged, effectively forming
a cone ca. 2 1.5cm, each individual cone subglo-
The species epithet refers to the whorled phyllotaxis bose or broadly ovoid, 46 35mm, yellow turn-
of the leaf-like cladodes. ing brown. Microsporophylls helically arranged,
imbricate, subpeltate, near abaxial base bearing two
Vernacular names yellow pollen sacs. Seed cones terminal or sublat-
eral, from a large, scaly bud, ovoid to cylindrical,
Japanese umbrella pine; koya-maki (Japanese) truncate, often irregular, 3.511 2.55cm, matur-
ing in second year, closed at first, often exuding
Description white resin, bract portion of bract-scale complex
brown, ovuliferous scale portion green, entire cone
Trees to 4045m tall, evergreen, monoecious; trunk turning brown with spreading scales. Bract-scale
monopodial, columnar, up to 3(3.5?) m d.b.h. Bark complexes helically arranged on a stout axis, more
thin and scaly, becoming fissured on trunk, exfo- or less dolabriform, up to 3.5 3.5cm; bracts and
liating in long, thin strips and flakes; inner bark ovuliferous scales proximally fused, with ovulifer-
reddish brown; outer bark dull brown. Branches ous scales exceeding the truncate bracts and these
spreading horizontally or occasionally ascending, terminating in a thick ridge with a central, ligulate,
forming a pyramidal crown in young trees and a 24mm long, thin, caducous bract tip; outer sur-
narrowly conical or cylindrical crown in mature face of ovuliferous scale portion deeply grooved,
trees. Foliage consisting of cladodes (needles) glabrous; cones when falling often partially disin-
in pseudowhorls on shortened terminal sections tegrated. Seeds inverted, 813 68mm (including
of long shoots, the density of this foliage depen- 2 wings), compressed; base truncate; apex acute;
dent on shoot elongation. Long shoots glabrous, surface smooth, brown with white hilum.
Taxonomic notes Conservation
For a detailed discussion, with references, of the seg- Being a component species in mixed conifer-
regation of the Japanese umbrella pine as the sole angiosperm forests where Chamaecyparis obtusa is
extant member of a distinct family Sciadopityaceae commonly the dominant species, past logging and
(and why it is not a yew [Taxus] nor a pine [Pinus]), subsequent conversion to managed or planted forest,
I refer to A Monograph of Cupressaceae and or other land uses, has restricted the occurrence of
Sciadopitys (Farjon, 2005a). Sciadopitys in many regions. Most of the remaining
stands in Honshu are now only small remnant popu-
Distribution lations and larger stands in more or less undisturbed
978 forest are confined to Shikoku and Kyushu. Past
Japan: Honshu (Aichi, Fukushima, Hiroshima, reduction has been inferred to have been ca. 20% and
Kyoto, Nagano, Nara, Okayama, Wakayama), the decline has now slowed down or possibly ceased.
Shikoku, Kyushu (Myazaki). IUCN: NT
TDWG codes: 38 JAP-HN JAP-KY JAP-SH
Uses
Ecology
The durable wood of this species is used for construc-
Locally common in mixed conifer-angiosperm for- tion purposes and to a limited extent for boat build-
ests, with Chamaecyparis obtusa, C. pisifera, Tsuga ing and certain kinds of furniture. The fibrous bark
sieboldii, Abies firma, Pinus parviflora, and angio- was formerly used for caulking boats (oakum). Its
sperms such as Aesculus turbinata, Magnolia obo- most valuable commercial use is undoubtedly in the
vata, Acanthopanax spp., Cercidiphyllum japonicum, horticultural industry, as it is a popular planted tree
and Acer rufinerve. In dense forest usually only ferns, in Japan and (more limited) in Europe and the USA.
mosses and liverworts grow under the trees, but in Apparently it was not introduced in China, as were
some localities shade tolerant shrubs and small a few other Japanese conifers such as Cryptomeria
trees such as Ilex sugeroki and Skimmia japonica can japonica. The Dutch, who were the only Europeans
thrive. Sciadopitys occurs as solitary trees as well as in allowed to trade with Japan, brought it to Jawa early
small groves or more or less pure stands, reflecting in the 19th century; it arrived successfully in England
events of recruitment and subsequent successional and the Netherlands in 1861. A small number of cul-
stages in the forest. This conifer is most commonly tivars with dwarfed growth, variegated foliage and/
found in rocky, cool and moist ravines and valleys in or pendulous branches has been developed but these
mountainous areas, at altitudes between 200m and are rarely planted. Some of these seem to scarcely dif-
1700m a.s.l. fer from the species.
Sequoia Endl., Syn. Conif.: 197. 1847 (nom. cons.). Conservation of this generic
name was unnecessary (Straub et al. in Taxon 57 (2): 646 (2008). Type: Sequoia
sempervirens (D. Don) Endl. [Taxodium sempervirens D. Don] (Cupressaceae).
Gigantabies J. Nelson, Pinaceae: 77. 1866. (nom. thin strips, reddish brown to grey. Branches spread-
illeg.). According to Straub et al. in Taxon 57 (2): 646 ing, curved down except in top of tree, in old trees
(2008) this name was not validly published (Art. 32, with many reiterations, with numerous spreading
Art. 33.9) but it has been listed as validly published foliage branches forming a conical or pyramidal, in
in Index Nominum Genericorum (ING). Type: large trees more rounded crown. Foliage branches
Gigantabies taxifolia J. Nelson [Sequoia sempervirens usually spreading horizontally and flattened, lat- 979
(D. Don) Endl.]. eral (ultimate) branchlets 312cm long, covered
with decurrent leaf bases, green turning reddish
Named after Sequoya (ca. 17601843), a Cherokee brown, semi-deciduous. Leaves alternate, decurrent,
chief who invented an alphabet for his people. on shaded shoots pectinate by a twist of lower free
part, not twisted but curved on most fertile shoots,
Description mostly linear, flattened, up to 25 3mm (usually ca.
20mm long but variable on the same shoot from
See the species description. base to middle to apex), straight or slightly curved;
margins entire; apex acute or acuminate, primarily
Distribution hypostomatic, with stomata adaxially in two broad
bands separated by a narrow midrib; leaf colour lus-
As for the species. trous green or dull grey-green to glaucous green, the
stomatal bands whitish or glaucous. Pollen cones on
the same branches as seed cones, subterminal or ter-
Sequoia sempervirens (D. Don) Endl., Syn. Conif.: minal from buds in leaf axils, solitary, ca. 5 4mm;
198. 1847. Taxodium sempervirens D. Don, in microsporophylls 1015, helically arranged, peltate,
Lambert, Descr. Pinus 2: [24], t. 7, f. 1. 1824. Type: with erose-denticulate margins, bearing 23 abaxial,
USA: California, Santa Cruz Co., Santa Cruz Mts., globose pollen sacs. Seed cones terminal on short
[California: Santa Cruz Mr. Menzies on back of branchlets, 1530 1318mm, maturing to purplish
sheet], A. Menzies s.n. (holotype BM). Fig. 319, 320 brown and finally (reddish) brown. Bract-scale com-
plexes 1825, helically arranged, parting when cone
Etymology matures except distal 23, peltate; distal part irregu-
larly diamond-shaped, 610 45mm, rugose, with
The species name means evergreen and was given to a central depression and transverse ridge in which
contrast this conifer with the deciduous Taxodium the bract tip forms a 1.52.5mm long process until
distichum, then thought to belong in the same genus. it erodes; proximal part narrowing to a pedicellate
base. Seeds 60100 per cone, irregularly shaped, 56
Vernacular names 34mm, flattened, with angular margins, reddish
brown, often with dark spots; wings 2, rudimen-
Redwood, Coast Redwood, California Redwood tary, forming a 1mm wide continuous margin. This
species is the only known hexaploid (chromosome
Description count 2n = 6x = 66) in conifers.
Giant Sequoia, Sequoia, Bigtree, Sierra Redwood; The discovery of this tree by Europeans was an event
Wellingtonia (United Kingdom) well publicised in the popular press of the time, due
to its sensational size which at first caused disbelief summers, with mean annual precipitation between
among more cautious scientists. Although first seen 9001400mm, but with high year-to-year variation.
by Albert Kellogg of San Francisco in 1852, specimens Temperature in winter is mild, with light frosts but
reached John Lindley of London late in 1853, and he occasional extremes, and warm, occasionally hot, in
first described it (hurriedly) as Wellingtonia gigantea summer. Sequoiadendron giganteum is well adapted
in the popular Gardeners Chronicle (of which he to low-intensity forest fires (extremely thick bark)
was editor) on Christmas Eve of that year (Ornduff and resists windfall exceptionally well; its wood is
in Aune, 1994). Kellogg, however, had an American rot-resistant. As a result its longevity ranges from
hero (Washington), not a British one, in mind for the 20003000(3200) years.
biggest tree on earth. The furious and highly nation-
982 alistic pathos that ensued, not only in the popular Conservation
press, but even in botany, is now amusing to read,
especially when we realise that Wellingtonia was, as Although nearly all groves are on public land, enjoy-
later homonym, illegitimate and that Washingtonia ing various levels of protection, the species is under
soon became the well-known name of a palm genus, IUCN criteria considered Endangered due to past
causing its use for the Giant Sequoia to be officially decline caused by exploitation, but particularly
rejected. because decline continues for other reasons caused
by past mismanagement in protected areas. Present
Distribution problems include fire risks, largely due to (past)
management practices which tended to benefit its
USA: California, Sierra Nevada (Calaveras, Fresno, coniferous competitors (especially Abies) rather
Madera, Mariposa, Placer, Tulare & Tuolumne Co.), than the target species, and which have greatly accu-
in ca. 67 disjunct groves. mulated the fuel load for future fires to burn more
TDWG codes: 76 CAL devastatingly. There is a considerable literature on
the conservation aspects of this species; for a compi-
Ecology lation see Aune (1994).
IUCN: EN [B2ab (ii, iii, v)]
This species is forming groves of a few to over
20,000 individuals in the Mixed Conifer Forest Uses
belt on the western slopes of the Sierra Nevada.
It is mixed with other conifers: Abies concolor, A. Since its discovery by Europeans in the mid-19th
magnifica, Calocedrus decurrens, Pinus lamber- century, exploitation during the latter half of that
tiana, P. ponderosa, P. jeffreyi, Pseudotsuga menzie- century and into the next was considerable. The
sii, Taxus brevifolia, and with fewer broad-leaved trees, though of high lumber quality and rot-resis-
trees: Quercus kelloggii, Q. chrysolepis, Cornus nut- tant, often shattered on impact of the giant boles.
tallii, Alnus rhombifolia, Salix scoulerana, Acer mac- What wood could be used was put into mainly
rophyllum, and shrubs: Castanopsis sempervirens, building applications, and many larger houses in San
Ceanothus cordulatus, C. parvifolius, C. integerrimus, Francisco and the Bay Area were built of its timber.
Arctostaphylos patula, etc. The relatively narrow alti- No commercial exploitation of wild groves occurs at
tudinal belt, (830)14002150(2700) m a.s.l., and present, and most of these were protected for their
the scattered concentration of groves, which tend scenic and scientific values many years ago. The
to become smaller and further apart going north, giant trees are a major international tourist attrac-
indicate rather narrow climatic and soil conditions tion in California. The Giant Sequoia is also highly
that are optimal in its natural habitat. Most groves regarded as an ornamental tree in parks and gardens
are on granite-based residual and alluvial soils, of large homes and, being easily propagated from
some on glacial outwash, and mildly acidic; best seed, is sold by many tree nurseries. Several cultivars
growth is on deep, well-drained sandy loams with have been named and are in the trade. The species
available ground water, the latter appears to be an also has potential as a managed-forest tree for tim-
important limiting factor. The climate is humid, ber production, but has found few applications in
with mostly autumn rain and winter snow, and dry commercial forestry thus far.
Sundacarpus (J. T. Buchholz & N. E. Gray) C. N. Page, Notes Roy. Bot. Gard.
Edinburgh 45: 378. 1989. Podocarpus LHr. ex Pers. sect. Sundacarpus
J. T. Buchholz & N. E. Gray, J. Arnold Arbor. 29: 57. 1948. Type: Sundacarpus
amarus (Blume) C. N. Page [Podocarpus amarus Blume] (Podocarpaceae).
Description
From Sunda, a name for the Indonesian archipel-
ago (minus the Moluccas and New Guinea); Greek Dioecious evergreen trees to 60m tall, with an erect
karpos = fruit. trunk to 1.5m diam., large trees with 35m tall but-
tresses. Bark smooth, dark brown weathering black- 983
Description ish grey, in large trees breaking into numerous square
plates; inner bark red-brown. Branches of trees in
See the species description. forest well above a clear bole, ascending or erect, then
spreading; foliage branches more or less pendulous.
Distribution Foliage buds small, globose, with imbricate, rounded
scales. Leaves on seedlings and juvenile plants dis-
As for the species. tinctly rostrate (abrubtly narrowed to an elongated
apex), 36(9) times as long as wide. Adult type
leaves usually narrower, broadly linear, 515cm long,
Sundacarpus amarus (Blume) C. N. Page, 615mm wide, petiolate and with a tapering base,
Notes Roy. Bot. Gard. Edinburgh 45: 378. 1989. with parallel or more or less undulating margins,
Podocarpus amarus Blume, Enum. Pl. Javae 1: 88. spreading pectinately or assugent, straight or irregu-
1827; Nageia amara (Blume) F. Muell., Select Pl., larly curved, with a central groove over the midvein
ed. 2: 138. 1876; Stachycarpus amarus (Blume) adaxially and a raised midrib abaxially, usually acu-
Gaussen, Trav. Lab. Forest. Toulouse T. 2, 1 (2, 20): minate at apex, dark green on the upper (grooved)
105. 1974 (nom. inval., Art. 33.2); Prumnopitys side, light green below. Stomata restricted to abaxial
amara (Blume) de Laub., Blumea 24 (1): 190. 1978. side (leaves hypostomatic), in two broad bands of
Type: Indonesia: Jawa, [locality not stated], leg. ign. numerous densely set lines. Pollen cones on stalks
s.n. (holotype L). Fig. 323, 324, 325 axillary to foliage leaves or subtended by decidu-
ous scale leaves, sometimes terminal and solitary,
Etymology more often 37 on a stalk, short cylindrical when
immature, elongating to 1530mm, 2.54mm wide;
The species epithet amaragiven by Blume under microsporophylls triangular-trullate, usually keeled,
Podocarpus (= amarus) means bitter and refers with serrate or entire margins and two longitudinally
to the taste of the fruit i.e. the fleshy epimatium dehiscing pollen sacs. Seed cones solitary or with 25
around the seed. on leafless pedunculate shoots 14cm long, borne
subterminally on 35mm long scaly dwarf shoots,
Vernacular names with distal scales with abortive ovules spreading out
or recurving and not enlarged or fused, with sterile
Black pine; choopoola (Queensland); sitobu scales falling off. Growing seeds covered by a green,
(Sumatera); ki merak, ki pait (Jawa); sempilau then orange to red and finally glaucous purple, fleshy
(Sabah); pasuig (Philippines) and numerous local but firm epimatium, becoming nearly spherical with
vernacular names given in Flora Malesiana, ser. 1, 10 a small protruding apex, 2230mm diam. Seed
(3): 387 (1988). proper globular, ca. 20mm diam., smooth with an
indistinct ridge. Cotyledons in 3 fused pairs.
Distribution Syzygium, and the conifer Phyllocladus hypophyl-
lus. At the highest altitudes it occurs in mossy forest
Malesia: from Sumatera and the Philippines to and becomes stunted. In Queensland S. amarus is a
New Guinea, New Ireland and New Britain (not in constituent of complex notophyll vineforest, a type
Peninsular Malaysia and in Borneo only in Sabah); of tropical rainforest with abundant lianas, where it
Australia: NE coastal Queensland. occurs with Podocarpus dispermus.
TDWG codes: 42 BOR-SB JAW LSI-ET LSI-LS MOL
PHI SUL SUM 43 BIS NWG-IJ NWG-PN 50 QLD-QU Conservation
Ecology IUCN: LC
984
Sundacarpus amarus is a tree of tropical evergreen Uses
rainforests, holding out in primary forest as scat-
tered tall and sometimes giant emergents and re- Sundacarpus amarus is a timber tree of sometimes
establishing in secondary forest. It is rare at sea level, very large dimensions and is logged often together
but becomes common from 500 to 2200m a.s.l. with other podocarps. It is sometimes distinguished
and has been found to 3000m on Mt. Kinabalu in in the timber trade as black podocarp. Its wood is
Sabah. At lower to middle elevations it is associ- sawn and used for construction (e.g. government
ated with conifers such as Agathis spp., Dacrycarpus buildings in remote mountain areas of Papua New
spp., Falcatifolium gruezoi, and Dacrydium spp.; Guinea), carpentry and joinery, as well as furni-
angiosperms are Cryptocaria pomatia, Dysoxylum, ture making. It is not well known in the trade as it
Macaranga, Ficus and many other tree species. It is still being confused with other podocarpaceous
grows often in latosols derived from andesite, basalt, species or treated with other species in the genus
or granite, rarely in sandy soils or in marshes. In Prumnopitys (PROSEA, 1993). Outside some collec-
New Guinea it is very common in montane forests tions in botanic gardens it is not known in horticul-
with Castanopsis, Nothofagus, Calophyllum, Pasania, ture or as a planted forestry tree.
Taiwania Hayata, J. Linn. Soc., Bot. 37: 330. 1906. Type: Taiwania cryptomerioides
Hayata (Cupressaceae).
Named after the island of Taiwan, from where it was dal crown in young trees, in old trees domed or flat-
first described. topped. Foliage branchlets (sub)pendulous, in young
trees with long subulate leaves, in older trees covered
Description with short, appressed scale leaves, persistent. Leaves
alternate to helically arranged, short decurrent,
See the species description. dimorphic with tree age; juvenile leaves persistent for
30 years or longer (trees ca. 15m tall), falcate-subu- 985
Distribution late, (5)1024 1.53.5mm, widest near base, with
free part spreading, bilaterally flattened, keeled on
As for the species. both faces, nearly straight or curved forward, acute-
pungent, amphistomatic, glaucous-green; adult
leaves short lanceolate-trullate, 36(7) 1.23mm;
Taiwania cryptomerioides Hayata, J. Linn. Soc., margins entire; apex acute to obtuse, incurved, free
Bot. 37: 330. 1906. Type: Taiwan: Nantou Co., or nearly appressed; leaves amphistomatic, lustrous
Chiayi-Nantou border, Wusungkengshan, (dark) green with whitish stomata. Pollen cones in
[ad pedem montis Morrison], N. Konishi s.n. terminal clusters of (2)35(7) on branchlets with
(lectotype TI). Fig. 326, 327, 328 scale leaves, ovoid-globose, 23mm long; microspo-
rophylls numerous, helically arranged, peltate, with
Taiwania flousiana Gaussen, Trav. Lab. Forest. (2)3(4) abaxial pollen sacs. Seed cones terminal,
Toulouse T. 1 (3, 2): 6. 1939. solitary, maturing in one season to ellipsoid to cylin-
Taiwania yunnanensis Koidz., Acta Phytotax. drical brown cones (9)1220(25) 611mm. Cone
Geobot. 11 (2): 138. 1942. scales (=bracts) gradually transformed from scale
leaves below, (12)1420(25), broadly obdeltoid to
Etymology obtrullate, 610 58mm; proximal portion cune-
ate, yellowish brown or reddish brown; distal por-
The species epithet means resembling Cryptomeria tion dull brown; apex mucronate. Seeds 1430 per
and refers to the type of leaves prevalent in young cone (12 per scale), ovate-oblong, flat, 4 23mm
trees. (without wings), light brown or tan. Wings 2, partly
overlapping, surrounding the seed, thin filamentous,
Vernacular names hyaline, 12mm wide.
Coffin tree, Taiwania, Taiwan cedar; tai wan shan Taxonomic notes
(Chinese)
The very disjunct occurrence in Yunnan and
Description Myanmar [Burma] in the west and Taiwan in the east
(the locus classicus) seems to have been the primary
Trees to 6065(70) m tall, evergreen, monoecious; reason why botanists have made taxonomic distinc-
trunk monopodial, straight, up to 34m diam. tions. A careful and critical comparison of types and
above buttressed base. Bark on large trees relatively many other specimens revealed no consistently dif-
thin, exfoliating in thin strips and chips, becom- ferent morphological characters (Farjon, 2005a).
ing fissured, reddish brown or brown weathering The recently discovered population in Viet Nam fits
grey. Branches spreading or assurgent, lower foliage in well with material from the earlier known loca-
branches pendulous, forming a conical or pyrami- tions; here it was observed that reversal to juvenile
type leaves can occur in lower parts of the crown in sheltered side valleys. It can attain an age of 1600+
of mature trees. Transition from juvenile cryp- (probably over 2000) years and belongs ecologically
tomerioid foliage leaves to adult sequoiadendroid to those conifers which through longevity and canopy
leaves occurs at an advanced age of trees, even in emergence survive all other forest trees, waiting for
the wild. episodal forest disturbance (most likely fire but pos-
sibly also landslides) to regenerate. The forest soils are
Distribution yellow and red acidic derivatives of granitic or meta-
morphic rocks. The climate is strongly influenced
China: NW Yunnan, SE Xizang [Tibet]; NE Myanmar by monsoons, with annual precipitation exceed-
[Burma]; Taiwan: Nantou District; Viet Nam: ing 4000mm in China, but at about 3000mm in
986 Lao Cai, Van Ban District. [Other reported localities Viet Nam. Reports of this species from other areas in
in China are here considered to be based on intro- China (Fu & Jin, 1992; Flora of China 4, 1999) do not
duced trees]. refer to this type of extremely wet monsoon forest and
TDWG codes: 36 CHC-YN CHT 38 TAI 41 MYA VIE it is very unlikely that Taiwania is indigenous there.
Ecology Conservation
Taiwania cryptomerioides is a conifer of montane In Yunnan many stands of old growth Taiwania are
forests at altitudes from 1750m to 2900m a.s.l. In still liable to be exploited; the establishment of more
Taiwan it grows in the cool temperate coniferous reserves for this species is urgent. In Taiwan the
forest belt with Chamaecyparis obtusa var. formo- establishment of Yushan National Park in 1984 pro-
sana and C. formosensis as codominant species and tected several natural stands of trees, but many had
more scattered occurrence of Calocedrus formo- been felled by that time. Plantation forestry using
sana, Cunninghamia konishii, Picea morrisonicola, this species is limited in Taiwan and more extensive
Pseudotsuga sinensis, Taxus wallichiana, and Tsuga in Guizhou and Hunan in mainland China; it will
chinensis. Angiosperm trees are common but scat- take many years before these are trees suitable for
tered, e.g. Castanopsis, Quercus and Trochodendron harvest as the species is (beyond sapling and pole
aralioides, while shrubs such as Camellia brevistyla, stages) slow growing. In Viet Nam, the small pop-
Eurya, Rhododendron, and Vaccinium are more abun- ulation is Critically Endangered (CR), because the
dant and the bamboo Yushania niitakayamensis can forest remnants in which it still occurs are acutely
cover large areas. In Yunnan and accross the border threatened by deliberately set fires in the area; it is
in Myanmar [Burma] it occurs in the mixed conifer- estimated that perhaps as much as 80% of its habi-
ous forest with Abies forrestii, Picea brachytyla, Larix tat has already been destroyed. The situation in
potaninii, Pseudotsuga sinensis, and Tsuga dumosa, Myanmar is unknown at present.
and with Cephalotaxus fortunei, Taxus wallichiana IUCN: VU (A2c, d)
and Torreya grandis var. yunnanensis in the under-
storey. All the trees are densely hung with the lichen Uses
Usnea longissima and mosses and leafy liverworts
cover trunks and branches. Some angiosperm trees The wood of Taiwania is very durable and valued
are mixed in and become more abundant at lower for timber; notable is the (past?) use of it for coffin
altitudes, e.g. Acer, Castanopsis, Lithocarpus, Quercus, making. In Viet Nam it was observed that the local
Magnolia, Schima, and Sorbus, and in the understorey HMong mountain people use it together with that
Rhododendron and many other shrubs are abundant. of Fokienia hodginsii for their houses, in particular
In Viet Nam it is scattered in remnants of montane for roof planks. Some of this wood is beautifully
evergreen forest dominated by Fagaceae, Lauraceae marked with red and pale yellow annual rings (late
and some Magnoliaceae, with only one other large and early wood) and prized for furniture. This tree
conifer, Fokienia hodginsii, common. In these forests has been planted in China outside its natural habi-
Taiwania is an emergent, usually forming small groves tat for much longer than the century that has passed
since its botanical description in 1906. It was intro-
duced to Japan, Europe, North America, and New
Zealand as an ornamental tree, where it grows well
outside in temperate regions with none or only light
frosts in winter and sufficient moisture. Despite its
attractive form and foliage as a young tree, it has
remained rare and is almost restricted to arboreta
and botanic gardens.
987
Taxodium Rich., Ann. Mus. Natl. Hist. Nat. (Paris) 16: 298. 1810. Type: Taxodium
distichum (L.) Rich. [Cupressus disticha L.] (Cupressaceae).
Cupressus disticha L. var. nutans Aiton, Hort. Kew. Pneumatophores or knees uncommon, if present
3: 372. 1789; Taxodium distichum (L.) Rich. subsp. more rounded (not conical). Short shoots assurgent
nutans (Aiton) E. Murray, Kalmia 12: 25. 1982. or erect. Leaves 58 ranked, largely appressed but
with free apices, not twisted at base, short acicular
990 Vernacular names (subulate) to narrowly lanceolate, 310mm long,
keeled, acute or pungent.
Bald-cypress, Cypress, Southern Cypress, Swamp
Cypress, Red-cypress, Yellow-cypress, White- Distribution
cypress, Gulf Cypress
SE USA: Coastal Plain from Virginia to E Texas, gen-
Description erally not as far inland as T. distichum var. distichum.
TDWG codes: 77 TEX 78 ALA FLA GEO LOU MSI
Pneumatophores or knees common and numer- NCA SCA VRG
ous, conical, 1(4) m tall. Short shoots spreading, but
with occasionally assurgent or nearly erect branch- Ecology
lets with short subulate leaves on the same tree, or
with intermediate forms. Linear, flattened leaves pec- As for the species, but more often in swamps and
tinately arranged, twisted near base, 1015(22)mm near lakes more distant from rivers.
long, straight or slightly curved, obtuse or mucronate.
Conservation
Distribution
IUCN: LC
As for the species.
992
figure 304. Pseudo
tsuga japonica foliage
and young seed cone
figure 339. Tetraclinis articulata seed cones figure 340. Thuja koraiensis in Korea
(photo M. Gardner) (photo Y. S. Kim)
figure 343. Thuja plicata foliage and seed
cones
figure 354. Tsuga mertensiana var. mertensiana figure 355. Widdringtonia cedarbergensis
foliage and seed cones seed cones
figure 359. Wollemia nobilis bud and foliage
Conservation Etymology
1008
Exploitation has reduced the population in many The species epithet means contorted, but it is
areas of China, but the species is still wide-spread. unclear what William Griffith had in mind when he
Logging in Viet Nam may have had a bigger impact chose this epithet.
as populations there are small; it may have reduced
the species to some extent to the less accessible Vernacular names
places. Exploitation of its bark and foliage for chemi-
cal/medical purposes has caused this species to be West Himalayan yew; mi ye hong dou shan
listed on CITES Appendix II. The species has poor (Chinese); postil (Kashmir); thuner (Hindi)
regeneration, hence any exploitation would deplete
the population. Harvesting can only be sustained Description
from plantations, as indiscriminately stripping trees
of bark and foliage may kill them. In China such Shrubs or trees to 15(20) m tall; trunk of trees
plantations have been established in several places. monopodial, short, or multi-stemmed, to 3m d.b.h.
IUCN: EN (A2d) but usually hollow in large specimens, with reiter-
ated stems inside the old bole; tree bases often read-
Uses ily coppicing. Bark thin, exfoliating in large irregular
flakes, reddish brown. Branches numerous, ascend-
The wood of this species is used in China in con- ing to erect, then spreading, frequently reiterating,
struction, cooperage, for the making of furniture, forming a rounded or pyramidal crown. Foliage
and for wood carving and turning. In Viet Nam it is branchlets irregularly alternate, slender, terete, with
also used for irrigation paddles in rice fields. Extracts fine grooves alongside decurrent leaf bases, green
of many parts of the plant (roots, wood, bark and turning yellowish brown to grey. Terminal buds
leaves) are used in traditional Chinese medicine, small, ovoid, with imbricate, closely appressed,
while in modern times the pharmaceutical industry acute, brown scales persisting at base of new branch-
became much interested in the anti-tumor proper- lets; lateral buds frequent, dormant. Leaves on lateral
ties of some of its alkaloids (in particular taxanes, shoots more or less arranged in V-formation, spaced
drug name: Taxol), which appear to be present in well apart, linear, 1.54(4.5)cm long, only slightly
all species in low but varying concentrations in bark twisted at short petiolate base, nearly straight or
and leaves. The seeds contain oils which are also straight, 1.52.5mm wide, with parallel, revolute
extracted, but treatment is required to neutralize the margins and a cuspidate apex. Midrib on adaxial
poisonous alkaloids. In horticulture it has been used (upper) side raised, ca. 0.2mm wide, nearly contin-
in bonsai and to a limited extent as a garden shrub. uous to apex, on abaxial side flat, ca. 0.4mm wide
It is doubtful whether this species is in cultivation and continuous to apex; leaf colour lustrous dark
outside China and Viet Nam. green above, two pale yellowish green bands below.
Stomata in two bands on abaxial side, densely and
randomly distributed. Pollen cones axillary, solitary,
forming rows on either side of fertile shoots, ovoid,
68mm diam., short pedunculate with dry, yellow- usually under canopy as a secondary layer shrub
ish green or brown, increasingly larger bracts at base; or tree to 12m tall, at elevations between 1800 and
axis slightly elongating at anthesis. Microsporophylls 3100(3400) m a.s.l.
614, peltate, each with 49 partly fused pollen sacs
on underside, pinkish turning brown. Seed-bearing Conservation
structures axillary, solitary, sessile, with tiny trian-
gular scales covering a very small dwarf shoot and Exploited intensively in recent years by extraction
a single terminal ovule except micropyle. Aril green of foliage and bark for its alkaloid chemical com-
at first, covering lower half of seed, swelling to suc- pounds (taxanes) which yield paclitaxel (drug name:
culent red and overtopping seed, leaving its apex Taxol) and similar chemicals, from which a drug
free, cup-like, 912mm long, 79mm wide. Seeds against ovarian and breast cancer in humans is pro- 1009
oblong, slightly flattened, obtusely ridged on two duced, this species has come under threat of extinc-
sides distally, with a small mucro, 67 45mm, tion. It was originally described from a limited area
green turning brown or black. in the Tibetan Himalayas (Xizang, China) but is
now understood to have a much wider distribution,
Taxonomic notes encompassing all of the western range of the species
commonly known as T. wallichiana. The latter spe-
Taxus fuana Nan Li & R. R. Mill was considered by cies is listed on CITES Appendix II; this should now
its authors to be distinct from T. wallichiana and T. apply to both species.
yunnanensis; the latter species is now commonly IUCN: EN (A2a, c, d)
treated as synonymous with T. wallichiana. Recent
morphometric analysis of the Himalayan yews con- Uses
firmed this separation (Mller et al., 2007). Spjut
(2007) concluded that the long forgotten name Recent discovery of anti-cancer effective com-
Taxus contorta Griff. has to replace T. fuana because pounds in the alkaloid taxanes of this and other spe-
they are the same taxon. In Flora of China 4: 8990 cies of Taxus has led to the exploitation of foliage
(1999), three species are keyed out with characters and bark which effectively kills the shrub or tree. In
that in part have to be observed from living (or at India, Nepal, and Pakistan this species is commonly
least not herbarium pressed) foliage; two of these known as T. wallichiana; a more detailed account of
involve the Himalayan and Sino-Himalayan taxa T. uses is given under that species.
contorta and T. wallichiana. Re-examination by me
of the herbarium specimens held at the Herbarium at
Kew (K) and the Natural History Museum, London Taxus cuspidata Siebold & Zucc., Abh. Math.-Phys.
(BM) under T. wallichiana confirmed the separation Cl. Knigl. Bayer. Akad. Wiss. 4 (3): 232. 1846.
into two taxa. The (fuzzy) dividing line seems to lie
in eastern Nepal, with all specimens collected west Etymology
from there assignable to T. contorta.
The species epithet refers to the leaves terminating
Distribution in a small cusp.
Ecology Description
In Nepal and Xizang [Tibet] this species occurs in Shrubs or trees to 25m tall; trunk to 1m (1.5m)
pine forests or in mixed conifer-angiosperm forests, d.b.h., often branching low. Branches long and
spreading or ascending, forming a broadly rounded, Ecology
usually compact crown. Bark thin, shallowly fis-
sured, exfoliating in strips or flakes, reddish brown. Taxus cuspidata occurs sparsely in mixed coni-
Foliage branches numerous, horizontally spread- fer and conifer-deciduous broad-leaved forests in
ing, erecto-patent or nearly erect, smooth with fine lowland to lower montane altitudes from 100m
grooves along the decurrent leaf bases, green turn- to 1600m a.s.l. In NE China it occurs in coni-
ing pale orange-brown or reddish brown. Terminal fer forest with Abies nephrolepis, Picea jezoensis,
buds ovate, with imbricate but partly free, broadly Pinus koraiensis, and Larix gmelinii var. olgensis,
elliptical or ovate, weakly keeled, green with hyaline in Sakhalin Island and northern Japan with Abies
margins, persistent at base of branchlets and turn- sachalinensis, Picea glehnii, P. jezoensis, Larix
1010 ing brown. Leaves distichously spreading, directed kaempferi, and various angiosperm trees e.g. Acer
more or less horizontally or often upwards (forming spp., Betula spp., Populus maximowiczii, Juglans
a V-shape) from lateral shoots, or sometimes more mandshurica, Sorbus aucuparia, Ulmus spp., and
radially spreading, linear, (1.1)1.53(3.5)cm long, Kalopanax ricinifolium. Further south in Japan it
straight or curved near base, 2.23.5mm wide; leaf is common in the understory of woods with Acer
base petiolate, twisted, decurrent; margins mostly spp., Ulmus davidiana var. japonica, Tilia japonica,
parallel, revolute, abruptly narrowing to a mucronate Juglans ailanthifolia, Quercus mongolica var. gros-
or sometimes acute apex. Midrib on adaxial (upper) sesserata, and many other species of trees. It grows
side slightly elevated, 0.2mm wide and continuous on a variety of soils derived from granitic, schis-
to apex, nearly flat and 0.30.4mm wide on abaxial tose or serpentine base rocks. The variety nana
side. Leaf colour lustrous dark green above, deep is mostly found growing on rocky sea coasts but
green with two pale yellowish green bands below. may also occur on exposed rock outcrops in the
Stomata on abaxial side in two bands separated by a interior.
midrib and margins, numerous, arranged in irregu-
lar lines. Pollen cones axillary, arranged in rows on Conservation
second year branchlets, sessile, globose, 3.54.5mm
diam., at base with imbricate, broadly ovate scales This species has been listed on CITES Appendix II
increasing in size, turning from pale yellow (on mar- in connection with the exploitation of its foliage for
gins) to pale reddish brown. Microsporophylls 915 the extraction of chemicals active as an anti-cancer
per cone, peltate, each bearing 58 obovate-cuneate, drug. This exploitation was localized and has not
pale yellow pollen sacs. Seed bearing structures axil- resulted in significant decline throughout the wide
lary, solitary, with numerous ovate or broadly ovate range of this species.
scales at base, partly enclosing a single terminal
ovule. Arils with a few pairs of scales at base, obovate, Uses
1013mm long, 911mm wide, with a round open-
ing showing the seed, succulent, deep scarlet or pur- The wood of this yew is used in China in construc-
plish red. Seeds broadly ovoid or trichonous-ovoid, tion, cooperage, for the making of furniture and
56mm long, 44.5mm wide, with 3(4) obtuse for wood carving and turning. In Japan it is prized
ridges below acute or mucronate apex, lustrous for interior finish, household furniture, utensils,
chestnut brown when ripe. marquetry, wood turning and sculpture; on a more
industrial scale it is used to make pencils. The Ainu
Distribution people of Hokkaido and Sakhalin made their bows,
as well as the scabbards of hunting knives, from its
Russian Far East: Kuril Is., Sakhalin, Primorye; wood. The heartwood yields a brown or red dye.
China: Heilongjiang, Jilin, Liaoning, Shaanxi; North Extracts of many parts of the plant (roots, wood,
Korea; South Korea; Japan. bark and leaves) are used in traditional Chinese
TDWG codes: 31 KUR PRM SAK 36 CHM-HJ CHM-JL medicine (treating diabetes) while in modern
CHM-LN CHN-SA 38 JAP-HK JAP-HN JAP-KY times the pharmaceutical industry became much
JAP-SH KOR-NK KOR-SK interested in the anti-cancer properties of its alka-
loids (taxanes, drug name: Taxol), which appear Description
to be present in all species in low but varying con-
centrations. The seeds contain oils which are also Large shrubs or trees. Leaves 1.53.5cm long, more
extracted, but treatment is required to neutralize or less distichously arranged.
the poisonous alkaloids. In horticulture it has been
so popular for a long time that the species is now Distribution
rare in the wild, but it is grown nearly everywhere
in the urbanised areas of Japan. Numerous culti- As for the species.
vars exist, and the species is used in bonsai culture.
Japanese yew was introduced to Europe (England) Conservation
in 1855 by Robert Fortune and more cultivars have 1011
been developed here as well as in the USA. A hybrid IUCN: LC
between T. baccata and T. cuspidata (Taxus media
Rehd.) originated in the USA around 1900 and has
given rise to further cultivars. Taxus cuspidata Siebold & Zucc. var. nana hort. ex
Rehd., in Bailey, Cycl. Amer. Hort. 4: 1773. 1902.
2 varieties are recognized: Taxus umbraculifera (Siebold ex Endl.) Lawson var.
nana (Rehd.) Spjut, J. Bot. Res. Inst. Texas 1 (1):
281. 2007. Type: Japan: Honshu, Hyogo Pref.,
Taxus cuspidata Siebold & Zucc. var. cuspidata. Mt. Hyonosen, G. Murata 44671 (neotype A).
Taxus baccata L. subsp. cuspidata (Siebold & Zucc.)
Pilg. in Engler, Pflanzenr. IV.5 [18]: 112. 1903. Taxus caespitosa Nakai var. angustifolia Spjut, J. Bot.
Type: Japan: [locality not stated], P. F. von Siebold Res. Inst. Texas 1 (1): 268. 2007.
s.n. (lectotype not designated, syntype K).
Fig. 337, 338 Vernacular names
Taxus wallichiana Zucc., Abh. Math.-Phys. Cl. Shrubs or trees to 30m tall; trunk of trees monopo-
Knigl. Bayer. Akad. Wiss. 3: 803, t. 5. 1843. Taxus dial, to 1.5m d.b.h. Bark thin, exfoliating in strips or
1016 baccata L. subsp. wallichiana (Zucc.) Pilg., in irregular flakes, variously coloured, reddish or pur-
Engler, Pflanzenr. IV.5 [18]: 112. 1903. Type: India: plish brown or grey. Branches numerous, ascend-
(eastern), N. Wallich s.n. [ex Herb. Zuccarini] ing to erect, then spreading or drooping, frequently
(lectotype M). reiterating, forming a spreading, rounded or pyra-
midal crown. Foliage branchlets irregularly alter-
Cephalotaxus celebica Warb., Monsunia 1: 194. 1900; nate, slender, terete, with fine grooves alongside
Taxus celebica (Warb.) H. L. Li, Woody Fl. Taiwan: decurrent leaf bases, green turning orange-brown
34. 1963. to purplish brown. Terminal buds small, ovoid,
Taxus yunnanensis W. C. Cheng & L. K. Fu, Acta with imbricate, closely appressed, rounded, dark
Phytotax. Sin. 13 (4): 86. 1975; Taxus wallichiana brown scales mostly early deciduous at base of new
Zucc. var. yunnanensis (W. C. Cheng & L. K. Fu) branchlets; lateral buds frequent, dormant. Leaves
C. T. Kuan, Fl. Sichuan. 2: 215. 1983; Taxus chinen- on lateral shoots more or less distichous, sometimes
sis (Pilg.) Rehd. var. yunnanensis (W. C. Cheng & slightly overlapping, linear to lanceolate, (1)1.5
L. K. Fu) L. K. Fu, Vasc. Pl. Hengduan Mount. 1: 214. 3.5(4.5)cm long, twisted at short petiolate or nearly
1993. sessile base, usually falcate, (1.5)24(5)mm wide,
Taxus sumatrana (Miq.) de Laub., Kalikasan 7 (2): thin and soft; margins flat or slightly revolute, more
151. 1978; Cephalotaxus sumatrana Miq., Fl. Ned. or less gradually tapering to a cuspidate apex or
Ind. [Fl. Ind. Batavae] 2 (7): 1076. 1859. more abruptly ending in a mucronate apex. Midrib
Taxus phytonii Spjut, J. Bot. Res. Inst. Texas 1 (1): 237. on adaxial (upper) side raised, 0.20.3mm wide,
2007. nearly continuous to apex, on abaxial side flat, ca.
Taxus contorta Griff. var. mucronata Spjut, J. Bot. 0.5mm wide and continuous to apex, usually with-
Res. Inst. Texas 1 (1): 260. 2007. out papillae; leaf colour lustrous dark green above,
Taxus florinii Spjut, J. Bot. Res. Inst. Texas 1 (1): 222. pale green with two pale yellowish bands below.
2007. Stomata in two bands on abaxial side, densely and
Taxus suffnessii Spjut, J. Bot. Res. Inst. Texas 1 (1): randomly distributed. Pollen cones axillary, soli-
226. 2007. tary, forming rows on either side of fertile shoots,
Taxus obscura Spjut, J. Bot. Res. Inst. Texas 1 (1): 235. ovoid, 56mm long, 34mm wide, short peduncu-
2007. late with dry, yellowish green to pale brown, increas-
ingly larger bracts at base; axis slightly elongating
Etymology at anthesis. Microsporophylls 814, peltate, each
with 46(8) partly fused pollen sacs on underside,
This species was named in honour of Nathaniel creamy white or pale yellow. Seed-bearing structures
Wallich (17861854), a Danish botanist who amassed axillary, solitary or sometimes in pairs, distributed
the famous Wallich Herbarium for the East India on underside in distal part of foliage branchlets, ses-
Company, now kept at Kew. sile, with tiny triangular scales covering a very small
dwarf shoot and a single terminal ovule except the
micropyle. Aril green at first, covering lower half of
seed, swelling to succulent red or orange (often more
or less translucent) and overtopping seed, leaving its
apex free, cup-like, 1013mm long, 710mm wide. Myanmar [Burma]; Viet Nam; Malesia: Philippines,
Seeds ovoid or obovoid, slightly flattened, with two Sulawesi, Sumatera.
obtuse ridges and a small mucro, 58 3.55mm, TDWG codes: 36 CHC-SC CHC-YN CHT 40 ASS-ME
green turning brown or black. ASS-NA EHM-AP EHM-BH EHM-DJ EHM-SI NEP 41
MYA VIE 42 PHI SUL SUM
Taxonomic notes
Ecology
In most of the literature referring to Taxus in the
Himalayas only one species, T. wallichiana, is recog- Taxus wallichiana is a small to large understorey
nized for the entire mountain chain [often referred or lower canopy tree in montane, temperate, warm
to as T. baccata subsp. wallichiana (Zucc.) Pilg.]. temperate, and tropical submontane to high mon- 1017
More recently, all of the populations occurring from tane forest, both angiosperm and conifer dominated,
central Nepal westward to northern Pakistan and deciduous or evergreen, or in mixed forests. In open
Afghanistan have been assigned to the relatively situations on rocky slopes and cliffs it usually forms
recently described species T. fuana Nan Li & R. R. a large, broadly spreading shrub. Elevation ranges
Mill, which was first known from the Himalayas of from 900m to 3700m a.s.l. and soils are mostly
SW Xizang [Tibet]. This species has turned out to derived from silicate-bearing rocks, i.e. acidic to
be synonymous with T. contorta Griff., a much ear- neutral. Like the European T. baccata it is easily
lier name, which has therefore priority and must be dispersed by birds and can germinate quickly in
used instead. It is quite distinct from both T. wal- large numbers on suitable sites. It has a very long
lichiana and from T. baccata, which does not reach life-span (theoretically indestructible) and sprouts
further east than N Iran. Two other species, T. chi- from stumps as well as roots. Taxus wallichiana
nensis and T. mairei, were included in T. wallichiana occurs in pure stands of limited extent or mixed in
as varieties in Flora of China 4 (1999), but here it the understorey of Quercus, Abies, Cedrus deodara,
is preferred to treat these as (not very) distinct spe- and Picea, or in mixed conifer forest. In Viet Nam
cies. In Malesia, the species Taxus celebica and T. T. wallichiana has been found growing in submon-
sumatrana, although considered distinct in recent tane evergreen mixed forests associated with the
treatments (Farjon, 1998 [2001], several Floras, Spjut conifers Cephalotaxus mannii, Dacrycarpus imbri-
2007) do not differ consistently in their morpho- catus, Keteleeria evelyniana, Nageia wallichiana, and
logical characters from T. wallichiana; indeed, ear- Podocarpus neriifolius. It will form dense thickets
lier botanists usually identified the specimens from on exposed rocky slopes with little tree growth. In
the Philippines, Sumatera, and Sulawesi as that spe- the Philippines it occurs on high ridges and moun-
cies. In some of these accounts, T. celebica and T. tain summits in mossy forest, or sometimes in rocky
sumatrana are reported to extend from Malesia into grass and scrubland.
Indochina and China, or even to Nepal, and there to
occur alongside T. wallichiana. Here these two spe- Conservation
cies are considered synonymous with T. wallichiana,
which therefore extends from Nepal to Sulawesi. In In certain regions of India in particular, harvesting
this place, it is not possible to comment in detail on of leaves and bark for extraction of alkaloid taxanes,
the recent work by Spjut (2007), other than to report used to produce the anti-cancer drug paclitaxel or
that comparison of the type specimens of his new similar chemicals, has led to a decline perhaps as
species from the region with specimens of T. wal- much as 90% in recent decades (Molur & Walker,
lichiana have caused me to conclude that they are 1998). Logging in Yunnan, China may have reduced
synonyms of that species. the rare populations in that country. Even though
its extent of occurrence (EEO) is very large and may
Distribution include areas where this kind of exploitation has not
occurred, its overall decline is inferred to have been
China: SE Xizang [Tibet], NW Yunnan, S Sichuan; in excess of 50%. Taxus wallichiana is listed under
E Nepal; Bhutan; India: Arunachal Pradesh, Assam; CITES Appendix II. It occurs in several protected
areas, e.g. the Sagarmatha National Park in Nepal. major reason for exploitation in recent years. The
Cultivation on a large scale in the context of phar- leaves yield similar chemicals in low concentra-
macology could reduce the pressure on wild grow- tions. Traditional medicine has made use of young
ing populations. shoots and leaves and sometimes of inner bark
IUCN: EN (A2a, c, d) for a long time in various potions, tinctures, and
pastes. The only non-toxic part of yews, the fleshy
Uses aril around the seed, is consumed by local inhab-
itants as jams. The inner bark also produces a red
The wood of Himalayan yew is durable and strong dye, often used in religious ceremonies by Brahmins
and is used for door frames, cabinet work and of Nepal (Singh, 2007). Since the exploitation of
1018 wood turning and wood inlaying, also for candle- Himalayan yew for its foliage containing taxanes
sticks, knife handles etc. Less refined products are has proved to be unsustainable, cultivation efforts
gates and fences, poles, struts and wattle and daub are being undertaken in the Himalayan foothills
in walls of rural buildings. The wood is also burnt and elsewhere in India; this involves both species T.
as incense in Nepal and parts of Tibet. The leaves contorta and T. wallichiana. These species, because
are toxic but can be given as fodder to goats if no of their similarity with European yew (T. baccata),
other foliage is available. The alkaloid compounds are rarely found in cultivation in Europe, but they
(taxanes) of the bark are a source for the anti-can- are used as ornamentals elsewhere, as in Baguio,
cer drug paclitaxel (Taxol) which has become a Philippines.
Tetraclinis Mast., J. Roy. Hort. Soc. London 14: 250. 1892. Type: Tetraclinis articulata
(Vahl) Mast. [Thuja articulata Vahl] (Cupressaceae).
Greek: tetra = four; kline = bed, couch; referring to long decurrent, adnate to shoot except apex, 1.68
the leaves in whorls of 4. 11.5mm (still longer on older branches, smallest on
ultimate branchlets), weakly dimorphic; facials lin-
Description ear, with broader, acute apex, partly covered proxi-
mally by the linear-spathulate, transversely convex
See the species description. laterals; apical margins denticulate; stomata in two
lateral rows or bands on facials, more sparsely on 1019
Distribution laterals; glands subapical, small, most prominent
on facials; abaxial leaf surface smooth, light green.
As for the species. Pollen cones terminal, solitary, ovoid-globose, ca.
4 2.5mm, reddish, maturing to yellowish brown
or brown; microsporophylls 812, decussate, pel-
Tetraclinis articulata (Vahl) Mast., J. Roy. Hort. tate, obtuse, bearing 4 abaxial pollen sacs. Seed
Soc. London 14: 250. 1892. Thuja articulata Vahl, cones solitary or sometimes clustered, terminal on
Symb. Bot. 2: 96. 1791. Type: Tunisia: Hammam-el- branchlets, maturing within one year to (sub)tetrag-
Lif, [legi copiose in montibus ad Hamamelis], leg. onal cones 1013 1217mm in size and becoming
ign. s.n. A (holotype C). Pl. 41, Fig. 339 first glaucous, then purplish brown to light brown.
Bract-scale complexes 4, decussate, nearly equal in
Etymology size, thick woody, ovate-cordate, 1012 812mm;
lower pair widest; upper pair with a truncate apex;
The species epithet refers to the seemingly articulate all abaxially smooth or rugose, with a subapical,
ultimate branchlets. small, recurved process (bract tip), adaxially striate,
with white seed scars near base, paler than exterior
Vernacular names parts. Seeds 46 per cone, conical-triangular, 45
3mm, brown, with dark spots; basal hilum whitish;
Sandarac; Thuya dAlgrie, Thuya de Barbarie, Bois wings 2, at oblique angle, large, ca. 10 6mm, reni-
de Citre; Arr (Algeria, Morocco); Alerce (Spain) form, thin menbranous, hyaline.
Description Distribution
Shrubs or small trees to 68(15) m tall, evergreen, N Algeria, Malta, Morocco, S Spain, N Tunisia, NE
monoecious; trunk monopodial or multistemmed, Libya (?); in Malta and Spain only tiny relict popula-
usually branching low or forked, up to 50cm diam, tions remain.
coppicing from base. Bark on large stems becoming TDWG codes: 12 SPA-SP 13 SIC-MA 20 ALG
rough, fissured, breaking into small plates, turning MOR-MO TUN
from light brown to grey. Branches long and slender,
more or less crooked angularly, forming a pyramidal Ecology
crown only in young, undisturbed plants, otherwise
irregular. Foliage branches articulate, branching Tetraclinis articulata occurs in the semi-arid to
alternately at various angles, ultimate branchlets winter rain parts of the Atlas Mountains and some
12mm thick, seemingly costate, persistent. Leaves scattered localities in the coastal hills near sea level;
decussate, appearing in whorls of 4 on thinnest there often on limestone, mixed with maquis. More
branchlets, obviously decussate on thicker branches, commonly in subclimax vegetation with Pistacia
1020
Plate 41. Tetraclinis articulata. 1. Habit of tree. 2. Branch with foliage and seed cones. 3. Branchlet with
pollen cones. 4. Pollen cone. 5, 6. Microsporophylls with open pollen sacs and pollen. 7. Immature seed
cones. 8, 9. Seed cones. 10. Seed cone scale. 11, 12. Seeds
lentiscus, Quercus ilex, and Juniperus phoeni- Uses
cea, locally into the zone with Cedrus atlantica,
on S-facing, warm and dry slopes to 1800m, on From the time of Phoenician and Roman coloni-
N-facing slopes to 1300m a.s.l. Woodland with this sation in North Africa to the present, the wood of
species is often severely degraded; however, this spe- Tetraclinis, and especially of the coppiced stools that
cies survives both fire and browsing by mans animals formed burrs (lupias), was prized and used for fur-
through its coppicing capacity, unless the pressure niture, cabinet making, and wood turning due to its
becomes too severe. Old coppice stools have very intricate markings and figuring from adventitious
dense wood and may be very ancient, but dating is growth. Today, only the larger trunks and stools with
very difficult. their contorted wood structure are valuable; most of
the lighter wood is used for fuel locally, especially 1021
Conservation in regions deficient in trees and with a poor rural
population. The resin (sandarac) is used as incense
The populations in Malta and southern Spain are in religious ceremonies and also forms a basis for
highly threatened. In North Africa, however, the varnishes and laquer work; Arabic tribes use(d) it
estimated area of occupancy is in excess of 1mil- for medicinal purposes. In cultivation it was for-
lion ha (Pardos & Pardos, 1997), with the largest merly propagated by the French colonial powers
populations in Algeria. Despite the capacity to cop- (Service dEau et Forts) in North Africa in attempts
pice, there are indications that increased browsing to extend its use as coppice wood in semi-arid areas.
pressure, especially by goats, prevent trees from It is uncommon as an amenity tree or garden orna-
regrowing in many areas (Gaussen, 1968, shows a mental and outdoor planting is restricted to coun-
photograph from Morocco to this effect). tries with a Mediterranean climate.
IUCN: LC
Thuja L., Sp. Pl. 2: 1002. 1753. Type: Thuja occidentalis L. (Cupressaceae).
Greek: thyia = resinous tree, also citrus tree. 1b. Mature seed cones (7)816(18) mm long.
Foliage sprays coarse, with small and larger
Description leaves; facial leaves usually glandular 2
2a. Seed cones with 68(10) scales. Pollen cones
Shrubs or small to large trees to 75m tall, evergreen, with 46microsporophylls T. occidentalis
monoecious; trunk monopodial or multistemmed; 2b. Seed cones with 812(14) scales. Pollen cones
bark fibrous, exfoliating in longitudinal flakes or with (6) 816microsporophylls 3
1022 strips. Branches spreading, curved down or assur- 3a. Seed cones with 812 scales, the two fertile
gent to ascending, forming a conical to pyramidal pairs of equal size, more or less truncate. Shrubs
crown; branching according to Massarts model. or small trees T. koraiensis
Foliage branches plagiotropic, branchlets alternate 3b. Seed cones with 812(14) scales, only one
to subdistichous, ultimate branchlets often more pair of equal size, obtuse or acute. Normally
numerous on the acroscopic side of lateral branch- large trees 4
lets, lateral branchlets compressed, persistent. 4a. Seed cones 1016(18)mm long; largest scales
Leaves decussate (semi-whorled), on whip shoots narrow, l/w ratio > 1.5. Ultimate branchlets on
long decurrent, on lateral branchlets dimorphic; foliage sprays more numerous on one (acro-
facials obtrullate-rhombic, smaller than laterals; lat- scopic) side of lateral branchlets, forming regu-
erals broadly falcate, spreading from apex of facials, lar tapering units T. plicata
conduplicate; margins entire; most leaves amphisto- 4b. Seed cones 712(14) cm long; largest scales
matic, but upper facials often epistomatic; stomata broad, l/w ratio < 1.5. Ultimate branchlets on
most numerous on lower surface of laterals; glands foliage sprays alternately arranged, forming
inconspicuous or absent. Pollen cones terminal, soli- irregular, more rounded units T. standishii
tary; microsporophylls 416, each with 24 abaxial
pollen sacs. Seed cones terminal, solitary or rarely
in pairs, maturing to ovoid-globose, narrowly ovoid Thuja koraiensis Nakai, Bot. Mag. (Tokyo) 33: 196.
or elliptical, partly opening cones. Bract-scale com- 1919. Type: North Korea: Hamkyongnam-do Prov.,
plexes 612(14), decussate; lowest pair reduced; Samsu-gun, Paek-san, V. L. Komarov 85 (lectotype
23 following pairs enlarged and spreading; upper LE). Fig. 340, 341, 342
pairs gradually smaller and narrower and apical pair
often connate. Seeds 410 per cone, flattened, with 2 Etymology
wings. Seedlings with 2cotyledons.
The species epithet refers to Korea, from where it
5 species. was first known to science.
Asia: disjunct in China: Chongqing (Dabashan), Korean Arbor-vitae; chao xian ya bai (Chinese);
Jilin (Changbai Shan); Korea; Japan. North America: nioi-nezuko (Japanese); nun chuk paek (Korean)
(disjunct) E Canada & USA; W Canada & USA
(coastal to N California). Description
Key to the species of Thuja Shrubs or small trees to 610m in their native habi-
tat; trunk often multistemmed, or monopodial,
1a. Mature seed cones 58mm long. Foliage sprays assurgent especially on slopes, to 30cm d.b.h. Bark
delicate, with very small eglandular leaves on stems thin, exfoliating in longitudinal flakes, red-
T. sutchuenensis dish brown or purplish brown. Branches spreading
to assurgent or ascending, forming a low, layering pumila, growing underneath its canopy; on lower
shrub or a pyramidal crown. Foliage branches pla- slopes it occurs as a small tree in taller conifer forests
giotropic; branchlets alternate to subdistichous; with Abies nephrolepis, Picea koraiensis, P. jezoensis,
third year shoots abruptly turning red-brown; lat- Pinus koraiensis, or P. sibirica; less often it grows in
eral branchlets compressed, with underside more clearings in mixed deciduous woodland in areas
flattened than convex upperside, persistent. Leaves with thin, rocky soil. It appears to avoid rocks of vol-
decussate, imbricate, on lateral branchlets dimor- canic origin and grows most abundantly on exposed,
phic; facials obtrullate-rhombic; laterals broadly granitic slopes and crags with acidic skeletal soil. The
falcate, spreading from apex of facials, conduplicate; climate is cool to cold temperate with high rainfall in
leaves variable in size from 110mm long on a single the mountains and prolonged winter snow cover.
branchlet system (facials smaller than laterals); mar- 1023
gins entire; apex acute to obtuse, of laterals incurved, Conservation
adnate; stomata most numerous on lower surface of
laterals in a slightly sunken band; glands small, often The extent of occurrence (EOO) calculated for this
visible on facials; leaf colour when newly emerged species, based on distribution data of herbarium
often glaucous, turning dark green; stomatal bands collections, is 48,091 km2 and the area of occupancy
white. Pollen cones subglobose, 23 23mm, pur- (AOO) is unknown. This small, shrubby tree is not
ple maturing to purplish black; microsporophylls exploited for timber, but may be cut for its foliage
610, peltate, each with 34 abaxial, yellow pol- used as incense. Most subpopulations are small,
len sacs. Seed cones solitary or sometimes in pairs, scattered and declining, and although detailed infor-
maturing in one year (2 seasons) to ovoid-globose; mation for North Korea is not available, it is now
opening cones 711 69mm with brown scales. thought that fewer than 1000mature plants exist in
Bract-scale complexes 812, decussate; lowest pair the wild. The only sub-population in China occurs
reduced, recurved; two following pairs enlarged to within a protected area; in South Korea it occurs in
56 26mm and spreading; upper pairs gradually Sorak N. P.
smaller and narrower, to 1mm wide, often connate; IUCN: VU [B2ab (iiv); C2a (i); D1]
larger scales abaxially striate or with longitudinal
grooves and a small, subapical bract tip, adaxially Uses
rugose, with two seed depressions. Seeds 810 per
cone, ellipsoid, 45 13mm, flattened, light brown; Because most plants in the wild are shrubs, this spe-
wings 2, marginal, 12mm wide, meeting with a cies is not useful for its wood. It was introduced to the
notch at both ends, thin menbranous, with minute USA by Ernest Wilson in 1917, from where it came
puberulence near the distal part. to Europe shortly after; however it remains relatively
rare in cultivation and is mostly restricted to arbo-
Distribution reta and botanic gardens. This is partly due to diffi-
culties of access of new material, as most populations
NE China: Jilin (Changbai Shan); North Korea occur in North Korea. Remarkably, in cultivation in
(Pyongannam-do, Pyonganpuk-do, Hamkyongnam- the UK, under conditions with ample moisture as in
do, Kangwon-do); South Korea (Kyongsanpuk-do, Hergest Croft Gardens, Herefordshire, some plants
Kangwon-do, Cholla-namdo). have grown into a tree much taller than any known
TDWG codes: 36 CHM-JL 38 KOR-NK KOR-SK in the wild in Korea; it shows no signs of being near
its limits at ca. 15m tall. This form is apparently based
Ecology on E. H. Wilsons collection No. 9244made on 12
October 1917 on Kongo-san at 1500m a.s.l. in South
Thuja koraiensis occurs on middle and upper slopes Korea. In the Netherlands, a very glaucous tree form
of mountains, at altitudes between 750m and 1950m is grown in the Gimborn Arboretum and University
a.s.l. Where slopes are exposed, it forms dense, low of Utrecht Botanic Garden. In NE China and Korea
thickets, but in sheltered ravines it can become a the shrubby form is used as an ornamental in gar-
small upright tree. It is often associated with Pinus dens and some public parks on a limited scale.
Thuja occidentalis L., Sp. Pl. 2: 1002. 1753. Type: 812 46mm, erect at maturity. Bract-scale
USA: [locality unknown], P. Kalm LINN 1136.1 complexes (6)8(10), decussate; lowest pair much
(lectotype LINN). reduced; higher pairs narrowly oblong; upper pair
connate or fused; two largest pairs spreading, ca. 8
Etymology 4mm, (ob)ovate-oblong, thin, obtuse or acute, with
a subapical bract tip 0.2mm, abaxially cinnamon or
The epithet means: from the west; Linnaeus named light brown, dull, adaxially light orange-brown, lus-
another species orientalis (from the east, i.e. China), trous, without apparent seed marks. Seeds 48 per
but this is now accommodated in a different genus. cone, usually only 4 (well) developed, 4 12mm,
flattened, light orange-brown; wings 2, marginal, of
1024 Vernacular names equal size and shape, surrounding seed but leaving
a notch at one or both ends, 11.5mm wide, thin
Northern White-cedar, White-cedar, Eastern White- membraneous, translucent.
cedar, Arbor-vitae, Swamp Cedar, Thuier cdre,
Cdre blanc Taxonomic notes
Uses
Dolophyllum Salisb., Quart. J. Sci. Lit. Arts 2 (4): 313. facials dolabriform or spathulate, obtuse; laterals
1817 (nom. rej.). Type: Dolophyllum dolabrata (L. f.) spreading from below their apex, bifacially flattened;
Salisb. [Thuja dolabrata L. f.]. margins entire; apex free, incurved, obtuse or acute;
the broad primary stomatal bands all on underside
Greek: opsis = aspect, appearance, i.e. appearing of plagiotropic shoots, on both sides on erect shoots;
1030 similar to the genus Thuja. leaf colour above lustrous dark green; stomatal
bands conspicuous, white, bordered by light green
Description margins. Pollen cones terminal, solitary, ovoid-
globose, 34mm long, reddish purple maturing to
See the species description. red-brown; microsporophylls 810, decussate, pel-
tate, rounded, bearing 35 abaxial pollen sacs. Seed
Distribution cones terminal on straight branchlets with small
but unmodified scale leaves, subglobose, markedly
As for the species. umbonate, 816(20)mm long. Bract-scale com-
plexes 68(10), decussate; lower and upper pairs
reduced; 4 larger pairs cuneate or obdeltoid, apically
Thujopsis dolabrata (Thunb. ex L. f.) Siebold & thickened, subapically reflexed with a large 23mm
Zucc., Fl. Japon. 2 (4): 34, tt. 119, 120. 1844. long, emerging and upturned bract tip, abaxially
rugose, dull brown to grey, adaxially striated, with
Etymology 25 seed scars at base, light brown. Seeds 1525 per
cone, narrowly ovate, irregularly compressed, 45
The species epithet, from Latin dolabra = hatchet or 2mm, greyish brown; wings 2, lateral, 11.5mm
mattock, refers to the shape of the facial leaves. wide, thin menbranous, yellowish.
Hiba Arbor-vitae; asunaro, asuhi, hiba (Japanese) Japan: Hokkaido, Honshu, Kyushu, Shikoku.
TDWG codes: 38 JAP-HK JAP-HN JAP-KY JAP-SH
Description
Ecology
Trees to 1520(30) m tall, sometimes a large shrub,
evergreen, monoecious; trunk monopodial or mul- This species is a constituent of conifer and conifer-
tistemmed, to 1m d.b.h.; branches frequently layer- angiosperm forests from lowland coastal areas to
ing. Bark on trunks with longitudinal plates, reddish montane sites in regions with a cool, moist climate.
brown with greyish strips of outer bark. Branches Its altitudinal range is 4001800(2100) m a.s.l. The
spreading or ascending to erect in (layering) shrubs, most commonly associated conifer is Tsuga diver-
forming pyramidal to broadly conical or domed sifolia, but it can grow in natural forests with sev-
crowns. Foliage branches in flat but irregular sprays, eral others, e.g. Chamaecyparis obtusa, C. pisifera,
branching alternately or divaricately; small branch- Cryptomeria japonica (locally), Thuja standishii,
lets variable in length, distinctly flattened, persistent. Sciadopitys verticillata (locally), Abies homolepis, A.
Leaves decussate, imbricate, dimorphic (on older, mariesii, and Pinus parviflora. Common angiosperm
thickening branches becoming monomorphic, trees are Betula ermanii, Fagus sieboldii, Quercus
decurrent and gradually widening with the growing mongolica var. grosseserrata, Pterocarya rhoifolia,
shoot), (1)38 15mm on flattened branchlets; Aesculus turbinata, and Cercidiphyllum japonicum.
Only occasionally does this species form more or less Distribution
pure stands, this is mainly the case with the northern
variety T. dolabrata var. hondae. Elsewhere, trees are Japan: S Honshu, Kyushu, Shikoku.
mostly scattered and small and must be extremely TDWG codes: 38 JAP-HN JAP-KY JAP-SH
shade tolerant, growing under canopy of Tsuga or
Fagus and competing with e.g. Ilex rugosa (Wilson, Conservation
1916). The variable disposition of foliage branches and
the development of stomatal zones on leaves indicate IUCN: LC
adaptability to changing light conditions; the capac-
ity of layering is probably another strategy to main-
tain sufficient light interception under an expanding Thujopsis dolabrata (Thunb. ex L. f.) Siebold & 1031
canopy, especially in earlier phases of growth. Zucc. var. hondae Makino, Bot. Mag. (Tokyo) 15:
104. 1901 [hondai]. Type: Japan: Honshu, Aomori
Uses Pref., Aomori, [Prov. Mutsu, near Awomori],
T. Makino s.n.(?) (holotype TI).
Hiba Arbor-vitae is an important forest tree in Japan
and is one of the five trees of Kiso (all are conifers) Description
originally preserved for imperial use. There are now
managed state forests with this species as the domi- Foliage relatively slender, with on average slightly
nant tree besides some imperial forests. Its light, soft, smaller leaves than in var. dolabrata. Seed cones
resinless and durable wood is used in construction, 1520mm long, bract-scale complexes less
for bridges, buildings, furniture, the wooden basis of umbonate.
laquer work, wood carving, etc. Younger trees of this
species are valued for their ornamental qualities due Distribution
to a striking contrast between upper lustrous green
and lower white banded leaf surfaces. Hiba Arbor- Japan: Hokkaido, N Honshu.
vitae is widely planted in Japan as well as in other TDWG codes: 38 JAP-HK JAP-HN
countries with a temperate maritime climate. Several
cultivars are known, most common are those with Conservation
variegated leaves and some dwarf forms with com-
pact growth and smaller leaves. The earliest of these IUCN: LC
were selected in Japan and introduced to Europe at
the same time as the species, around the middle of
the 19th century.
Description
Caryotaxus Henkel & W. Hochst., Syn. Nadelhlz.: Key to the species of Torreya
366. 1865. Type: Caryotaxus nucifera (L.) Henkel &
W. Hochst. [Taxus nucifera L.]; Foetataxus J. Nelson, 1a. Leaves (2)36(9)cm long. Branches spread-
Pinaceae: 167. 1866. Type: Foetataxus montana J. ing and drooping 2
Nelson (nom. illeg.) [Torreya taxifolia Arn.]; Tumion 1b. Leaves 1.23.8(4.5)cm long. Branches spread-
Raf. ex Greene, Pittonia 2: 193. 1891. Type: Tumion ing and ascending 3
1032 taxifolium (Arn.) Greene [Torreya taxifolia Arn.]. 2a. Leaves mostly drooping; stomatal bands
0.51mm wide; margins on that side of leaves
Named after John Torrey (17961873), American 0.50.7 mm wide. Ripe fleshy aril including
botanist. seed 1012 mm wide, pruinose to reddish
yellow T. jackii
Description 2b. Leaves pectinately arranged; stomatal bands
0.30.4 mm wide; margins 0.71 mm wide.
Dioecious or rarely monoecious evergreen trees or Ripe fleshy aril including seed to 25mm wide,
sometimes shrubs. Resin canals in leaves (1) and green with purplish streaks T. californica
seed arils. Bark smooth or flaking, becoming fis- 3a. Leaves straight or slightly down-curved, with a
sured. Leaves alternate (helically inserted), pecti- long cuspidate or spinose apex T. nucifera
nately arranged, flattened, linear-lanceolate, rigidly 3b. Leaves straight or distally (slightly) falcate, with
coriaceous, asperous, sharply acuminate or pungent, a short cuspidate apex 4
with a prominent midrib. Stomata in two separated 4a. Leaves (0.7)12.7( rarely to 4.5)cm long; sto-
bands on abaxial side. Pollen cones solitary in the axil matal bands 0.30.8mm wide. Ripe fleshy aril
of each leaf on fertile shoots, forming double rows including seed pale purplish brown or whitish
on the underside of plagiotropic foliage branchlets, pruinose T.grandis
subglobose; microsporophylls decussately arranged, 4b. Leaves 1.23.8(4.5) cm long; stomatal bands
tightly clustered, peltate, with 36 pendulous pol- 0.30.5 mm wide. Ripe fleshy aril including
len sacs forming a small umbel and containing sub- seed green streaked with purple, or slightly
spherical, arillate pollen. Seed-bearing structures whitish pruinose 5
axillary to foliage leaves, consisting of a miniaturized 5a. Midrib on the abaxial (lower) side of leaves
branching shoot with a few spirally inserted bracts 0.30.6 mm wide, leaf margins on that side
on the main and lateral axes and usually 23 sub- 0.51.2 mm wide, total leaf width 24 mm.
terminal, erect ovules of which only 1 develops into Ripe fleshy aril including seed pale green or
a large, hard seed with a strongly sclerified seed coat slightly whitish pruinose T. fargesii
and completely surrounded (except at the distal tip) 5b. Midrib on the abaxial side of leaves 0.60.8mm
by a fleshy aril which ripens in the second year and wide, leaf margins on that side 0.60.8 mm
becomes purplish green to bluish black. Seedlings wide, total leaf width 23.2mm. Ripe fleshy aril
with 4 short cotyledons. including seed dark green streaked with purple
T. taxifolia
6 species
Distribution
This taxon is considered threatened on account of Torreya grandis Fortune ex Lindl., Gard. Chron.
deforestation in many areas where it occurs (or has 1857: 788. 1857.
occurred in the recent past according to herbarium
collections made since the beginning of the 20th Etymology
century).
IUCN: EN (A2c, d) The species epithet means great and may refer to
the potentially large size of the arillous seed, espe-
cially in some cultivated forms.
Vernacular names Taxonomic notes
Chinese nutmeg tree; fei shu, yuan bian zhong Recently, a new species, T. parvifolia, was described
(Chinese) from Sichuan in China, the type specimens of which
are deposited in local Chinese herbaria. Its descrip-
Description tion, with an illustration, made comparison with
T. yunnanensis, but not with T. grandis or any other
Trees to 25m tall; trunk to 1(2) m d.b.h. Bark irregu- species. Its reported characters fit well within those
larly vertically fissured, sometimes flaking, yellow- given here and elsewhere (e.g. Flora of China 4, 1999)
ish grey or greyish brown, weathering to dark grey. for T. grandis var. grandis and it is therefore treated
1036 Branches spreading and ascending, forming a broad here as a synonym of that taxon. A fair number of
crown. Foliage branchlets slender, usually with oppo- varieties and forms have been described in this spe-
site branching, sometimes with more than 2 laterals cies; several of these are based on cultivated plants in
from a node, spreading horizontally at 4070 from China, which would probably be more appropriately
main axis in most cases, terete, with grooves along be termed cultivars. They are often based on charac-
margins of decurrent leaf bases, green turning yellow- ters of the arrillous seeds and some of these forms
ish green and then grey after first year. Buds at ends attain large sizes and are of economic importance
of branchlets very small; bud scales at lower node of in Chinese traditional medicine. Only one recently
previous year enlarged, broadly triangular, keeled, described variety has been recognized in Flora of
lustrous brown, deciduous thereafter. Leaves pecti- China 4: 95 (1999); its seeds are differently shaped
nately arranged, spreading at 6590 from shoot axis, and its leaves are longer than in other varieties. It is
linear to linear-lanceolate, (0.7)12.7(4.5)cm long, accepted here.
straight or distally falcate, 23.5(4)mm wide, with
a twisted, 0.51mm long petiole, abruptly widening Distribution
at base, tapering to a cuspidate apex, coriaceous, lus-
trous green on the adaxial (upper) side, with (or with- China: Anhui, Fujian, Guizhou, Guangdong, Henan,
out) an indistict midvein at least in lower half of blade. Hubei, Hunan, Jiangxi, Sichuan, Zhejiang.
Stomatal bands on abaxial side only, 0.30.8mm TDWG codes: 36 CHC-GZ CHC-HU CHC-SC
wide, light brown, separated by a 0.40.7mm wide, CHS-AH CHS-GD CHS-FJ CHS-HE CHS-HN CHS-JX
green midrib and bordered by 0.50.8mm wide, flat CHS-ZJ
or very narrowly revolute green margins; stomata
small, randomly distributed. Pollen cones axillary, Ecology
solitary, forming short rows on underside of lower
leaves of lateral branchlets, with several pairs of basal Torreya grandis is a constituent of the diverse mixed
bud scales in 4 rows, ca. 58 4.55mm before anthe- mesophytic forest (remnants) in Zhejiang, in which
sis, pale yellow; microsporophylls numerous, pel- broad-leaved trees (angiosperms) dominate but
tate, each with 45marginal, pendulous pollen sacs. several conifers can be present as associated trees.
Seed bearing structures axillary, solitary or in pairs, The altitudinal range of this species is between
sometimes grouped together on a branchlet, sessile, 200m and 1400m a.s.l. It is commonly found along
with rounded, keeled bract scales subtending seed. streams and/or in shady spots in the forest, but large
Ripe fleshy aril including seed smooth, pale purplish trees can reach the canopy and compete well with
brown or slightly whitish pruinose, obovoid-conical many other tree species. In secondary vegetation it
to ovoid, or ellipsoid to oblong-ellipsoid, 2040mm may persist for some time, but eventually disappears
long, 1225mm wide, mucronate at apex. Seed proper due to excessive growth of bamboo and other weedy
smooth or with 2 opposite longitudinal ridges. plants.
Uses Torreya grandis Fortune ex Lindl. var. jiulongsha-
nensis Z. Y. Li, Z. C. Tang & N. Kang, Bull. Bot. Res.
This species yields high quality wood used in build- Harbin 15 (3): 356. 1995. Type: China: Zhejiang,
ing houses, bridges (durability of the wood), and Suichang, Jiulong Shan, Z. Y. Li & Z. C. Tang 9009
furniture; some of it is also used to make utensils (holotype PE).
and for wood turning. The seeds are edible and pro-
duce oil that is extracted; the essential torreya oil is Description
extracted from the aril. The seeds proper (kernels)
are sold in Chinese drug shops as a remedy against Leaves 2.54.5cm long. Ripe fleshy aril including
coughs and can be eaten as nuts. This species is com- seed obovoid-conical.
mon in cultivation in China (especially in Zhejiang), 1037
where some forms with various shapes and sizes (up Distribution
to 5cm long) of the arillous seeds are treated as cul-
tivars and these are planted as fruit trees or shrubs China: S Zhejiang (Suichang Xian).
in special plantations. The Chinese nutmeg tree was TDWG codes: 36 CHS-ZJ
introduced to England by Robert Fortune in 1855,
but remains uncommon in cultivation in Europe Conservation
and elsewhere. Where it is grown it develops into a
shrubby habit, not a monopodial tree. IUCN: DD
Ecology
Torreya nucifera (L.) Siebold & Zucc., Abh. Math.-
Torreya jackii occurs in evergreen broad-leaved Phys. Cl. Knigl. Bayer. Akad. Wiss. 4 (3): 234.
forest, along streams, on steep slopes in shade or 1846. Taxus nucifera L., Sp. Pl. 2: 1040. 1753. Type:
in secondary vegetation usually near moisture. Its Illustration: [Fi, vulgo Kaja, Taxus nucifera] in
altitudinal range is between 120m and 1320m a.s.l. Kaempfer, Amoen. Exot. Fasc.: 814, 815. 1712.
It is a species of warm temperate to subtropical (lectotype). Fig. 349
regions affected by the SE monsoon with an annual
precipitation of 1350 to 1600mm and mean annual Torreya ascendens Nakai ex Uyeki, [Not. Pl. Lign.
temperature 1720 C, with absulute minimum of Sikoku 11] Sci. Rep. Matsuyama Agric. Coll. 10: 3.
10 C. It grows on acidic mountain yellow earth or 1953.
Etymology mucronate at apex. Seed proper smooth or with 2
opposite longitudinal ridges.
The species epithet (Latin nucis = nut) means
bearing nuts. Distribution
Description Ecology
1039
Shrubs or trees to 25m tall; trunk to 1(2) m d.b.h. Torreya nucifera is a woodland species occurring
Bark irregularly vertically fissured, sometimes flak- scattered in most types of mixed broad-leaved-
ing in narrow strips, yellowish grey or greyish conifer forest in the southern half of Japan. Wilson
brown, weathering to dark grey. Branches spread- (1916) mentioned a locality SW of Tokyo, where it is
ing wide, forming a broad crown. Foliage branchlets more abundant, growing with Abies firma on steep
slender, usually with opposite branching, sometimes slopes composed of shale on a hill ca. 500m high. In
with more than 2 laterals from a node, spreading other sites, including some islands in South Korea, it
horizontally at 4080 from main axis in most cases, holds out in secondary vegetation where it becomes
terete, with grooves along margins of decurrent leaf shrubby. Old-growth forest with ancient trees of T.
bases, yellowish green turning reddish to purplish nucifera in large numbers occurs in Koreas Bija-Rim
brown after first year. Buds at ends of branchlets Forest, Halla-san National Park, with trees estimated
small; bud scales at lower node of previous year to be 500800 years old. The altitudinal range is
enlarged, broadly triangular, keeled, lustrous brown, from near sea level to at least 1100m a.s.l. Besides
deciduous thereafter. Leaves pectinately arranged, Abies firma and Tsuga sieboldii, the most common
spreading at 7590 from shoot axis, linear to linear- conifers among which Torreya nucifera is found,
lanceolate, (1.5)23.5cm long, straight or slightly Chamaecyparis obtusa, Podocarpus macrophyl-
down-curved, 2.23.5(4)mm wide, with 12mm lus, Nageia nagi, Taxus cuspidata, and Sciadopitys
long, twisted petiole, abruptly widening at base, verticillata are often associated with this species.
tapering to a long cuspidate or spinose apex, coria- Locations dominated by conifers in the mixed meso-
ceous, lustrous dark green on adaxial (upper) side, phytic forests of warm temperate Japan are often on
smooth or with two parallel grooves at least in lower rocky S-facing slopes with poorly developed soils.
half of blade. Stomatal bands on abaxial side only, 0.3
0.4mm wide, yellowish, separated by a 0.50.8mm Conservation
wide, green midrib and bordered by 0.71mm wide,
flat and slightly raised green margins; stomata small, IUCN: LC
randomly distributed. Pollen cones axillary, solitary,
forming short rows on underside of lower leaves of Uses
lateral branchlets, with several pairs of basal bud
scales in 4 rows, ca. 57 45mm before anthesis, The wood of this species is valued in Japan for its
pale yellow; microsporophylls numerous, peltate, lustrous, pale brown leaf colour and its durability
each with 45marginal, pendulous pollen sacs. Seed especially in contact with water. It is used for furni-
bearing structures axillary, solitary or in pairs, some- ture and cabinet making, chests and boxes, Japanese
times grouped together on a branchlet, sessile, with chessmen, and formerly water buckets. The seeds are
rounded, keeled bract scales subtending seed. Ripe rich in edible oils and these as well as the arils are
aril including seed smooth, dark green ripening used in Japanese cuisine. This tree is widely planted
with purplish brown striation near apex, obovoid to near temples and in parks and gardens in Japan; all
ovoid, or ellipsoid, 2030mm long, 1318mm wide, trees N of Tokyo are thought to be originally planted
(Wilson, 1916) and some of these are champion trees of basal bud scales in 4 rows, ca. 56 4.55mm
much larger than most trees in the forests. In Europe before anthesis, pale yellow; microsporophylls
and North America it is an uncommon ornamental numerous, peltate, each with 46marginal, pendu-
shrub or tree in arboreta, botanic gardens, and occa- lous pollen sacs. Seed bearing structures axillary, in
sionally in private gardens. pairs, sometimes grouped together on a branchlet,
sessile, with rounded, keeled bract scales subtending
seed. Ripe aril including seed smooth, dark green
Torreya taxifolia Arn., Ann. Mag. Nat. Hist., ser. streaked with purple, obovoid or broadly ellipsoid,
1, 1: 130. 1838. Type: USA: Florida, Appalachicola 2535mm long, to 25mm wide, mucronate at apex.
River, J. Torrey s.n. (holotype K). Seed proper smooth or with 2 opposite longitudinal
1040 ridges.
Etymology
Distribution
The species epithet compares the leaves with those
of yew (Taxus). SE USA: NW Florida, SW Georgia, mainly along the
Appalachicola River, with one population 11 km W
Vernacular names of this river in Jackson Co., Florida.
TDWG codes: 78 FLA GEO
Florida nutmeg tree, Stinking cedar, Gopherwood
Ecology
Description
Torreya taxifolia occurs along limestone bluffs on
Small trees to 13(18) m tall; trunk to 80cm d.b.h. the Appalachicola River in a region with a warm
Bark irregularly vertically fissured, sometimes and humid climate, occasionally influenced in win-
flaking, pale brown or greyish brown, weathering ter by cold waves from the north that dip tempera-
to dark grey. Branches spreading and ascending, tures below the freezing point. It grows mostly in the
forming an open, broadly conical crown. Foliage shade of wooded ravines and steep, N-facing slopes
branchlets slender, usually with opposite branching, under canopy of Fagus grandifolia, Liriodendron
sometimes with more than 2 laterals from a node, tulipifera, Acer barbatum, Liquidambar styraciflua,
spreading horizontally at 4060 from the main Quercus alba, and occasionally pines (Pinus taeda,
axis in most cases, terete, with grooves along mar- P. glabra). Often these woods are hung with vines
gins of decurrent leaf bases, green turning yellow- (e.g. Smilax spp., Bignonia capreolata). Another rare
ish brown after first year. Buds at ends of branchlets conifer, Taxus floridana, occasionally grows with
very small; bud scales at lower node of previous year Torreya taxifolia. A capacity to resprout from the
enlarged, broadly triangular, keeled, lustrous brown, stem base probably accounts now for most of the
deciduous thereafter. Leaves pectinately arranged, surviving individuals in the wild.
spreading at 5080 from shoot axis, linear to linear-
lanceolate, 1.53.8cm long, straight or very slightly Conservation
falcate, 23.2mm wide, with a 0.51mm long,
twisted petiole, abruptly widening at base, tapering This rarest of the species of Torreya is nearly extinct
to a cuspidate apex, coriaceous, lustrous green on in the wild. It is almost restricted to a 64 km stretch
adaxial (upper) side, slightly convex at least in lower of the Appalachicola River, where it occurs spo-
half. Stomatal bands on abaxial side only, greyish radically in a specific limestone bluff habitat. The
white, 0.4mm wide, separated by a 0.60.8mm number of individuals has been drastically reduced
wide, green midrib and bordered by 0.60.8mm in recent decades, probably caused by a pathologi-
wide, flat or very narrowly revolute green margins; cal fungal disease (Ascomycota) of the stem and
stomata small, randomly distributed. Pollen cones leaves, first observed in 1962 and presumably alien
axillary, solitary, forming short rows on underside of to the region, having arrived in the late 1950s.
lower leaves of lateral branchlets, with several pairs Currently between 500 and 1000 individual plants
remain, many only as resprouts from stumps. At other trees migrated north again. It will be doomed
the moment in situ conservation is considered not a if that microclimate were to disappear with ongoing
viable option; therefore, an extensive ex situ conser- global warming. However, deliberately moving this
vation programme has been established. This proj- species north into habitats from which it would have
ect is collaborative between the Botanic Garden of disappeared naturally if it ever occurred there has
Smith College and Atlanta Botanical Gardens, USA. also been criticized (Schwartz, 2005/08).
The programme has involved the collection of over IUCN: CR (A2a, c, e)
6,000cuttings taken from plants throughout the
natural range of the species. Rooted plants have now Uses
been shipped to gardens throughout the world in
the hope that, if and when the natural environment The hard and durable wood has been used for fence 1041
has stabilised, material from these plants can assist posts locally before it became too rare and protected
the restoration of the species in its natural habitat. under State and Federal Law. The use for Christmas
Recently, assisted migration to areas in the south- trees has likewise ceased. It is rare in cultivation
ern Appalachian Mountains has been proposed, outside ex situ conservation programmes aiming at
based on the hypothesis that the historical range of reintroduction in the wild. This species appears to be
this species in lowland Florida is a refuge from the hardy in countries with mild winters (it apparently
last Ice Age (Barlow & Martin, 2008). During the can withstand occasional hard frosts). Growing
previous interglacials it may have occurred in these this tree in arboreta and botanic gardens should be
mountains (the same could be true for Taxus flori- encouraged, as it could help its conservation while
dana) but, for various reasons, in the current inter- being an attractive small conifer tree. It is apparently
glacial it missed the last bus and was marooned in commercially available from tree nurseries in South
the cool microclimate of its present location, while Carolina, USA.
Tsuga (Endl.) Carrire, Trait Gn. Conif.: 185. 1855. Type: Tsuga sieboldii Carrire
[Abies tsuga Siebold & Zucc.] (Pinaceae).
USA: Appalachian Mountains in Georgia, North Chinese hemlock; tie shan (Chinese)
Carolina, South Carolina, Tennessee and Virginia.
TDWG codes: 78 GEO NCA SCA TEN VRG
Description are red and yellow earth, or mountain podzols at
high elevations. The climate is cool temperate, moist
Trees to 4050 tall, d.b.h. to 1.52m; trunk mono- (annual precipitation 1000mm to 2000mm) or very
podial, often forked above 1/2 height. Bark on trunk wet (Taiwan). This species is widely distributed and
rough and scaly, with numerous brown-grey plates. occurs in the mixed mesophytic forest formation
Branches assurgent, spreading more horizontally (Wang, 1961) together with numerous broad-leaved
at ends; crown broad, conical, or flat topped in trees and several conifers; on the Southwestern
old trees. Branchlets pale yellowish brown, finely Plateau also in the montane coniferous forests with
grooved between slightly swollen, appressed, decur- Abies, Picea and other conifers.
rent and darker pulvini, with minute pubescence
1046 in grooves, soon glabrous. Vegetative buds ovoid- Uses
globose, 14mm long, not resinous, dark brown or
red-brown. Leaves mostly pectinate, but a few short Chinese hemlock is a valuable timber tree in China
leaves erect, (0.6)12(2.7)cm long, 1.83mm and has been logged extensively in many parts of
wide, (broad) linear, flattened, grooved above, with the country. The wood is hard and durable and used
near apical margins of young leaves denticulate, for construction, shingles for roofing, general car-
emarginate, sometimes entire at apex; stomata in pentry, and joinery. This species was introduced by
two white bands on underside (abaxial side) sepa- Ernest Wilson for the Veitch Nurseries in England
rated by a midrib, leaf colour green on upperside. in 1900, but it has remained uncommon in cultiva-
Pollen cones crowded near ends of branchlets, tion. No cultivars have been recorded. In plantation
35mm long, yellow, with purple tinge. Seed cones forestry, it is increasingly planted in the eastern USA
numerous, short pedunculate or sessile, ovoid- as a substitute for T. canadensis and T. caroliniana,
oblong when closed, subglobular or ovoid-oblong because it is resistant to an insect pest that adversely
when opened, 1.53(4?)cm long, 1.32.2(3.5?)cm affects the native species.
wide, light green when immature, ripening to light,
glossy brown. Seed scales nearly circular, convex, 3 varieties are recognized:
812(14) 810(12)mm; abaxial surface usually
striated, shining, with imprints of two lower scales,
glabrous; upper margin rounded, truncate or retuse; Tsuga chinensis (Franch.) E. Pritz. var. chinensis.
base short pedicellate. Bracts transversely rhom- Abies chinensis Franch., J. Bot. (Morot) 13 (8):
bic, with denticulate upper margin, 12(3)mm 259. 1899. Type: China: Chongqing Municipality,
long. Seeds ovoid-oblong or ovoid-triangular, 34 Chengkou Xian, Daba Shan, P. G. Farges 808
2mm, light brown; seed wings obliquely ovate, 67 (holotype P). Fig. 350, 351
3.5mm, light yellowish, transparent.
Tsuga formosana Hayata, Gard. Chron., ser. 3, 43:
Distribution 194. 1908; Tsuga chinensis (Franch.) E. Pritz. var. for-
mosana (Hayata) H. L. Li & H. Keng, Taiwania 5: 64.
S and N Central and SE China, SE Xizang [Tibet] (?); 1954.
Taiwan; N Viet Nam. Tsuga patens Downie, Notes Roy. Bot. Gard.
TDWG codes: 36 CHC-CQ CHC-GZ CHC-HU Edinburgh 14: 16. 1923; Tsuga chinensis (Franch.)
CHC-SC CHC-YN CHN-GS CHN-SA CHS-AH CHS-FJ E. Pritz. subsp. patens (Downie) E. Murray, Kalmia 12:
CHS-GD CHS-GX CHS-HE CHS-HN CHS-JX CHS-ZJ 26. 1982; Tsuga chinensis (Franch.) E. Pritz. var. patens
38 TAI 41 VIE (Downie) L. K. Fu & Nan Li, Novon 7 (3): 263. 1997.
Tsuga tchekiangensis Flous, Bull. Soc. Hist. Nat.
Ecology Toulouse 69: 414. 1936; Tsuga chinensis (Franch.)
E. Pritz. var. tchekiangensis (Flous) W. C. Cheng &
Tsuga chinensis and its varieties occur at altitudes L. K. Fu, Fl. Reipubl. Pop. Sin. 7: 119. 1978.
between (600)12003200(3500) m a.s.l. The soils
Tsuga chinensis (Franch.) E. Pritz. var. daibuensis Tsuga chinensis (Franch.) E. Pritz. var. robusta W.
S. S. Ying, Bull. Exp. Forest Natl. Taiwan Univ. 114: C. Cheng & L. K. Fu, Acta Phytotax. Sin. 13 (4): 83.
150. 1974. 1975. Type: China: Hubei, Yangtse Gorges, [?] Chen
et al. 2000 (holotype PE).
Description
Description
Seed cones ovoid-oblong when closed, more or less
subglobose when opened, 1.52.5cm long, 1.32.2cm Seed cones ovoid-oblong when closed, more or less
wide; seed scales nearly circular, short petiolate, 812 subglobose when opened, larger than in var. chinen-
810mm. sis (possibly up to 4 3.5cm); seed scales robust and
thick. 1047
Distribution
Distribution
As for the species.
China: Hubei (Badong Xian), Sichuan (Yalong
Conservation Valley).
TDWG codes: 36 CHC-HU CHC-SC
IUCN: LC
Conservation
Distribution Etymology
China: Chongqing, Gansu (Zhouqu), W Hubei, The species epithet refers to the the different lengths
Sichuan. of leaves on a shoot.
TDWG codes: 36 CHC-CQ CHC-HU CHC-SC
CHN-GS Vernacular names
Japan: N and Central Honshu, Kyushu, Shikoku. Tsuga dumosa (D. Don) Eichler, in Engler & Prantl,
TDWG codes: 38 JAP-HN JAP-KY JAP-SH Nat. Pflanzenfam. 2 (1): 80. 1887. Pinus dumosa
D. Don, in Lambert, Descr. Pinus 2: 55. 1824. Type:
Ecology not designated.
Tsuga diversifolia occurs in the mountains at alti- Abies yunnanensis Franch., J. Bot. (Morot) 13 (8):
tudes between 700m and 2000m a.s.l., on usually 258. 1899; Tsuga yunnanensis (Franch.) E. Pritz., Bot.
podzolic soils developed on volcanic or igneous Jahrb. Syst. 29: 217. 1900; Tsuga dumosa (D. Don)
rock. The climate is cool, with cold, snowy winters Eichler var. yunnanensis (Franch.) Silba, Phytologia
and abundant rainfall in summer (annual precipita- 68: 73. 1990.
tion 1000mm to 2500mm). It is in many areas the Tsuga dura Downie, Notes Roy. Bot. Gard. Edinburgh
most common tree species in mixed coniferous for- 14: 16. 1923; Tsuga yunnanensis (Franch.) E. Pritz.
ests, being very shade tolerant. Other common coni- subsp. dura (Downie) E. Murray, Kalmia 12: 26. 1982.
fers are Picea jezoensis, Abies homolepis, A. veitchii, Tsuga wardii Downie, Notes Roy. Bot. Gard.
A. mariesii (at high elevations), Larix kaempferi, Edinburgh 14: 17. 1923; Tsuga chinensis (Franch.)
E. Pritz. subsp. wardii (Downie) E. Murray, Kalmia purplish brown. Seed scales broadly ovate-elliptic,
12: 26. 1982. convex, 1114 912mm at mid-cone; abaxial sur-
Tsuga leptophylla Hand.-Mazz., Akad. Wiss. Wien, face smooth, finely striated, with imprints of over-
Math.-Naturwiss. Kl., Anz. 61: 83. 1924; Tsuga lapping scales, glabrous; upper margin rounded,
dumosa (D. Don) Eichler subsp. leptophylla (Hand.- entire, slightly recurved or straight; base petiolate,
Mazz.) E. Murray, Kalmia 12: 26. 1982. slightly auriculate. Bracts broadly angular-ovate, ca.
3mm long. Seeds ovoid-oblong, 34 22.5mm,
Etymology brown; seed wings obliquely ovate, 68 3.54mm,
light yellowish brown, transparent.
Dumosa is a Japanese name for Tsuga (but not
referring to this species). Distribution 1049
This species has a limited range separated into two or Western hemlock, Pacific hemlock
three disjunct areas. Deforestation and logging have
substantially reduced the area of occupancy (AOO) Description
of this species. A reassessment in 2010considered
that decline has slowed but not ceased in which case Trees to 6070m tall, d.b.h. to 1.52.5m; trunk
a suspected past reduction of 30% meets the crite- straight, often more or less buttressed at base (from
ria for Vulnerable under A2. originating on a nurse log). Bark on trunk rough
IUCN: VU (A2c, d) and scaly, fissured below, dark grey. Branches
spreading horizontally, drooping at ends; crown 1051
Uses in young trees conical, with drooping leader, in
old trees broad conical. Branchlets reddish brown
Forrests hemlock is a timber tree used for construc- above, pale yellowish below, turning grey in second
tion, aircraft, furniture and as props for mines. This year, finely ridged and grooved between small, dark
species is present in several arboreta in Europe and tipped, appressed, decurrent pulvini, lanate pubes-
North America, almost exclusively from early 20th cent with short brown hairs, mixed with fewer
century introductions made by the famous plant long whitish hairs, glabrescent after third year.
hunters of the time. These planted trees usually bear Vegetative buds ovoid-conical, 1.52mm long, not
a good crop of seed cones with viable seed and in resinous, densely pubescent, pale brown or red-
sufficiently wet places like western Scotlands arbo- dish brown. Leaves of unequal length, with longer
reta, they can be seen to self-propagate seedlings. leaves spreading sideways or radially, parted below
However, since this species is rarely the only one of the shoot, 0.72(2.3)cm long, 1.52mm wide,
its genus planted there, we cannot be sure, without (very) narrowly elliptic, flattened, grooved above;
specialized genetic research, that the seedlings are margins denticulate; apex obtuse or slightly emar-
pure T. forrestii. This creates a problem for the main- ginate; stomata in 2 white bands on abaxial (under)
tenance of the species in cultivation. If taken from side separated by a midrib; leaf colour dark green
cuttings, we base its propagation on a very narrow on upperside. Pollen cones subglobular, 35mm
genetic basis; from seeds we may introduce alien long, crimson or light red, yellow at anthesis.
genes into the stock of young trees. Ideally, coni- Seed cones numerous on all outer branchlets, on
fers like this are continually introduced from cor- 46mm long, pubescent peduncles, ovoid-oblong
rectly identified natural sources. Under the current when closed, subglobular with opened seed scales,
restrictive legislation pertaining to plant collecting 1.42.2(3)cm long, 1.52.5cm wide, light green
and international traffic of the same and its propa- or grey-green, occasionally tinged violet, ripening
gules, this option has become much more difficult to light brown, but unexposed part of seed scales
to realize. In the long term, ill-considered legislation turning dark brown. Seed scales suborbicular,
may seriously hamper the perpetuation of labori- ovoid-oblong or oblong, thin, opening wide, 613
ously assembled species collections in arboreta and 59mm at mid-cone; exposed part of abaxial sur-
botanic gardens. face striated or wrinkled, glabrous; upper margin
rounded or obtuse; base short pedicellate. Bracts
triangular, 24 mm long. Seeds ovoid-oblong,
Tsuga heterophylla (Raf.) Sarg., Silva N. Amer. 12: 23.5mm long, (light) brown; seed wings ovate
73, t. 605. 1899. Abies heterophylla Raf., Atlantic J. triangular, 410mm long, light yellowish brown,
1: 119. 1832. Type: not designated. Fig. 353 transparent.
Etymology Distribution
The species epithet describes the variable size of the W North America: along the coast from Alaska
leaves on a single shoot. to N California and in the Cascade Range, also in
the northern Rocky Mountains (mainly in British tree; in the wetter parts near the Atlantic coast it will
Columbia and Idaho). regenerate spontaneously. As an extremely shade tol-
TDWG codes: 70 ASK 71 ABT BRC 73 IDA MNT ORE erant conifer it is suitable as an under planted tree in
WAS 76 CAL deciduous broad-leaved forest; however, as it in turn
shades out every growth under it, this should not be
Ecology done in (semi) natural woodland where the indig-
enous flora is valued. Although frequently planted as
Tsuga heterophylla occurs from sea level to 600m a specimen tree in arboreta and parks, mainly within
a.s.l. along the Pacific coast, in the Rocky Mountains its natural range and in the British Isles, this species
it reaches to 1800m. It grows on a variety of soils is not much used in horticulture and few cultivars
1052 with an acid organic top layer (pH 3.55). The cli- have been raised.
mate is cool maritime along the coast, cold mon-
tane in the interior, the annual precipitation varies
between (500)9003800mm, decreasing towards Tsuga mertensiana (Bong.) Carrire, Trait Gn.
the interior. Dry summers limit its range in the Conif., ed. 2, 1: 250. 1867.
Rocky Mountains. It is highly sympatric with Picea
sitchensis in most of the range. It is extremely shade Etymology
tolerant, but has a shorter life span than Pseudotsuga
menziesii or Picea sitchensis. Close along the coast This species was named after Karl Heinrich Mertens
it may form occasionally pure stands, but more (17961830), whose plant collections from the
commonly it is an important constituent of the American Pacific coast were studied by A. H. von
(maritime) mesothermal coniferous forest. On the Bongard in St. Petersburg.
Olympic Peninsula of Washington it reaches maxi-
mum size, together with other giant conifers. Its Vernacular names
tolerance to shade allows it to grow up under the
canopy of other trees, but a thick moss layer usu- Mountain hemlock
ally prevents the light seeds from reaching the soil.
Instead, seeds germinate massively on fallen trees Description
(nurse logs), from where a few saplings are able to
send roots down into the soil; as a result T. hetero- Trees to 3040(45) m tall, d.b.h. to 11.5m; trunk
phylla often stands in rows (collonades) long after straight, but often curved or almost prostrate at tree
the nurse log has rotten away. limit. Bark deeply fissured in the lower part of the
trunk, dark reddish brown. Branches spreading or
Conservation assurgent; crown usually narrowly conical, but often
deformed by wind. Branchlets yellowish or orange-
IUCN: LC brown, turning grey in 24 years, with ridges end-
ing in decurrent pulvini with oval leaf scars, densely
Uses yellowish pubescent. Vegetative buds ovoid conical,
23 12mm, not resinous, brown. Leaves densely
Western hemlock is an important timber tree in the covering shoot, directed forward, ascending above
Pacific Northwest (USA) and W Canada. Unlike its shoot, (0.5)0.72(2.5)cm long, 11.5mm wide,
eastern sister species it is a fast grower even outper- short petiolate at base, linear, straight or curved,
forming Douglas fir. The timber is used for pilings, thick, 34 sided or with convex sides, shallowly
poles and railway sleepers and especially for con- grooved near base, obtuse or acutish at apex; sto-
struction. A large proportion of the annual harvest mata in several lines on all sides; leaf colour green
goes to the wood pulp industry for various applica- or glaucous green. Pollen cones pendulous, ca. 1cm
tions. This species has been introduced in Britain and long, at first purplish blue, then yellow with purple
other parts of NW Europe as a forestry plantation tinge. Seed cones mostly near top of tree, erect at
first, later mostly pendant, short pedunculate or Uses
sessile, ovoid-oblong to cylindrical, obtuse at apex,
(2)35.5(8.1)cm long, (1.1)1.52.5(3.3)cm wide This slow growing species produces moderately
with opened scales, purplish blue when young, rip- strong wood, but its use is limited due to environ-
ening to dark brown or light brown. Seed scales mental considerations. It is soft and close-grained,
4072(80?), obovate-cuneate, 1013(20) 710( with brown heartwood, sometimes pinkish, and
15)mm at mid-cone, spreading very wide or reflexed lighter sapwood; its uses are now restricted to car-
when mature, puberulent when immature, but soon pentry and some limited construction applications.
glabrous; upper margin rounded, entire; base short In its natural habitat it is much appreciated by hikers
pedicellate. Bracts ligulate-cuspidate, 47mm long, for its picturesque appearance on mountain ridges. It
visible with opened seed scales. Seeds cuneate, 35 also makes an excellent ornamental tree for gardens 1053
22.5mm, brown; seed wings oval-oblong, 47 with its dense foliage growing from long and short
33.5mm, light brown. shoots and naturally conical habit. Despite this, it
is uncommon in gardens and only a limited num-
Distribution ber of cultivars is known. Among these the blue or
glaucous leaf forms or selections are especially val-
Pacific Coast Region of NW North America: from ued. High altitude provenances may be susceptible
Alaska to California, in the Cascade Range and to late frost, but those from more coastal northern
Sierra Nevada, and isolated occurrences in the areas should not suffer from this kind of damage as
northern Rocky Mountains. much.
TDWG codes: 70 ASK 71 BRC 73 ORE WAS 76 CAL
NEV 2 subspecies and 2 varieties are recognized:
Ecology
Tsuga mertensiana (Bong.) Carrire subsp. merten-
Tsuga mertensiana is a subalpine species, occur- siana var. mertensiana. Pinus mertensiana Bong.,
ring from near sea level in Alaska to 1500m a.s.l. Mm. Acad. Imp. Sci. Saint-Ptersbourg, sr. 6,
along the coast; in the Cascade Range between Sci. Math. 2: 163. 1832; Hesperopeuce mertensiana
1200m and 2100m a.s.l., and in the Sierra Nevada (Bong.) Rydb., Bull. Torrey Bot. Club 39: 100. 1912.
(subsp. grandicona) between 1800m and 3350m Type: not designated. Fig. 354
a.s.l. It grows on a variety of non-alcareous acidic
soils, sometimes on peat, more commonly on mor Description
humus (pH 3.13.9). Subsp. mertensiana is restricted
to a climatic zone with high precipitation, in British Short shoots apparent. Leaves usually crowded
Columbia between 2000mm and 4000mm per and ascending above shoots. Seed cones (2)35.5
year, with long, snowy winters and short, cool sum- (6)cm long; seed scales 5072(80?) in number,
mers. Subsp. grandicona grows in a much drier cli- 1013 710mm, dark brown.
mate, but there primarily on high, N-facing slopes.
The species is a major component of the Mountain Distribution
hemlock-Subalpine fir forest, occurring in pure
stands or mixed with Abies lasiocarpa, locally also Pacific Coast Region of North America: from Alaska
with A. amabilis, Picea glauca, P. sitchensis, P. engel- to N California, also Rocky Mountains in interior
mannii (Rocky Mts.), Pinus spp., Tsuga heterophylla, British Columbia and Idaho.
Xanthocyparis nootkatensis, Juniperus occidentalis, TDWG codes: 70 ASK 71 BRC 73 IDA ORE WAS
and Betula papyrifera. 76 CAL
Conservation
IUCN: LC
Tsuga mertensiana (Bong.) Carrire subsp. mer- Tsuga mertensiana (Bong.) Carrire subsp.
tensiana (A. Henry) C. K. Schneid. var. jeffreyi, grandicona Farjon, Proc. Kon. Ned. Akad.
in Silva-Tarouca, Uns. Freil.-Nadelhlzer: 294. Wetensch., C, Bot. 91 (1): 39. 1988. Type: USA:
1913. Tsuga pattoniana (Balf.) Sncl. var. jef- California, Mono Co., Mammoth Lakes, Twin
freyi A. Henry, in Elwes & Henry, Trees Gr. Brit. Lakes, B. Willard 54 (holotype RM).
Ireland 2: 231. 1907; Tsuga jeffreyi (A. Henry)
A. Henry, Proc. Roy. Irish Acad. 34: 55. 1919; Description
Hesperotsuga jeffreyi (A. Henry) C. N. Page, Notes
Roy. Bot. Gard. Edinburgh 45: 389. 1989. Type: not Short shoots apparent. Leaves usually crowded and
designated. ascending above shoots. Seed cones 3.58.1cm long,
1054 1.93.3cm wide; seed scales (4052)(60?) in num-
Description ber, 1218(20) 1015mm, light brown.
Distribution Etymology
Canada: British Columbia (Vancouver Island). This species was named after Philipp Franz von
TDWG codes: 71 BRC Siebold (17961866), who studied the flora of Japan
when stationed there in the employ of the Dutch
Ecology East India Company.
IUCN: DD Description
Pachylepis Brongn., Ann. Sci. Nat. (Paris) 30: 189. Seeds usually numerous, flattened or angular, with
1833, non Less. (1832). Type: Pachylepis cupressoides or without 2 wings of unequal size. Seedlings with
(L.) Brongn. [Thuja cupressoides L.]; Parolinia Endl., 24cotyledons.
Gen. Pl. Suppl. 1: 1372. 1841, non Webb (1840). Type: 4 species
Thuja cupressoides L. [= Widdringtonia nodiflora (L.)
Powrie]. Distribution
1056
Named after Samuel Edward Widdrington (1787 Southern Africa: from the Cape to Malawi.
1856, formerly Cook), Commander of the Royal
Navy. Key to the species of Widdringtonia
Predominantly in cool and wet mountain fynbos, Willowmore Cypress, Willowmore Cedar; Baviaans
often in rocky outcrops and among boulders on sum- kloofseder, Sapreehout (Afrikaans)
mits, or in montane grassland often near streams,
and in canyon woodland (kloofbos), accompanied Description
by numerous fynbos genera (e.g. Leucadendron,
Leucospermum, Metrosideros, Protea, Restio) or Trees to 2025m tall, now rarely to 40m; trunk
Pteridium, Myrica pilulifera, and Poaceae, or form- monopodial, to 11.5m d.b.h. Bark on trunks fis-
ing pure stands. The altitude ranges from 100m to sured, exfoliating in angular plates. Branches
2590m a.s.l. Soils are nutrient-poor, acidic, derived spreading, contorted, forming a pyramidal crown in 1059
mostly from granite, quarzite or sandstone. The cli- young trees, more irregular in older trees. Ultimate
mate varies from Mediterranean in the Cape region foliage branchlets 510mm long, more or less quad-
to subtropical with summer rains and tropical mon- rangular in cross section, 1mm wide, persistent.
tane in Malawi. Leaves on ultimate branchlets appressed, triangular-
ovate to rhombic, ca. 1.5 1mm, increasing in size
Conservation on older branchlets to 10 4mm, keeled at base;
upper margins minutely denticulate; apex acute,
This widespread species, which is capable of cop- incurved, often reflexed on thicker branchlets and
picing after above-ground destruction (fire), is not on whip shoots; stomata on both sides in two mar-
threatened with extinction. ginal lines; leaf colour dull light green. Pollen cones
IUCN: LC oblong, 35 1.52mm; microsporophylls peltate,
coriaceous, narrowly triangular to ovate, bearing
Uses 34 abaxial pollen sacs near base. Seed cones clus-
tered on foliage branchlets, sometimes terminal on
No commercial uses have been recorded for this short shoots, maturing in two growing seasons to
species. It may be readily coppiced and is probably irregular-globose, 2030mm diam., serotinous but
used for firewood locally. Its only horticultural use eventually wide openening cones. Bract-scale com-
seems to be limited to plantings in botanic gardens; plexes 4, thick woody, oblong, max. 25 15 10mm,
under glass where frost occurs and outdoors in curved slightly inward; outer surface smooth or stri-
regions with a mild climate. It is suitable for plant- ate, strongly verrucose or tuberculate along margins,
ing in countries with a Mediterranean climate (and with a prominent subapical, 58mm long, thick,
is planted in California) and should be used more recurved boss enclosing bract apex, purplish brown
often, provided it has first been assessed and found or brown; inner surface with angular, 23mm
not likely to become invasive. long seed scars at base, yellow and lustrous brown.
Columella short, thick, angular, to 6mm tall. Seeds
1418(20?) per cone, ovoid-oblong, 56 34mm,
Widdringtonia schwarzii (Marloth) Mast., J. Linn. slightly flattened, curved towards cuspidate apex,
Soc., Bot. 37: 269. 1905. Callitris schwarzii Marloth, yellowish brown ripening blackish brown, with
Bot. Jahrb. Syst. 36: 206. 1905. Type: South Africa: dull lighter hilum at base; wings 2, on either side of
Cape Province, Willowmore Distr., Kougaberge, seed, exceeding 34mm beyond retuse seed apex,
R.Marloth 3614 (holotype SAM). unequal, thin, more or less translucent.
Etymology Distribution
This species was named after Herr E. Schwarz who South Africa: Eastern Cape Province, Willowmore
drew Marloths attention to this tree. District, Kougaberge, in canyons [kloofs] draining
into the Baviaanskloof and tributaries.
TDWG codes: 27 CPP-EC
Ecology Vernacular names
Widdringtonia schwarzii occurs in rocky ravines on Mulanje Cypress, Mulanje Cedar; Nkungudza
steep slopes or cliffs (krantzes or kranse) and in (Chichewa)
(dry) river beds of canyons between (600)900
1200m a.s.l. Trees can attain large size in rocky Description
streambeds of canyons, sheltered from veld fires,
where they can form small groves. The climate is Trees to 4050m tall; trunk monopodial, usually
Mediterranean with mostly winter rain; the dry, hot straight, to 11.5(2?) m d.b.h. Bark on trunks becom-
summers are somewhat tempered by the sheltered ing soft fibrous, to 24cm thick, fissured, brown-
1060 microclimate in deep, shady canyons, where deeper grey, exfoliating in long strips. Branches spreading
ground water remains available under stream beds. or ascending, forming a pyramidal, eventually irreg-
ular or flat-topped crown. Ultimate foliage branch-
Conservation lets (3)720mm long, more or less quadrangular in
cross section, 11.5mm wide, persistent. Leaves on
In the past, settlers have exploited this species, which ultimate sterile branchlets appressed or with some
is capable of growing to large trees, to near extinction free apices, ovate to rhombic, 1.53.5 11.5mm,
in all accessible areas. In recent years, new localities increasing in size on leading branchlets (whip
have been found in remote upper parts of canyons in shoots) to ca. 10 4mm, weakly keeled at base;
the Kouga Mountains, some of which harbour large upper margins minutely denticulate; apex obtuse to
trees. There is now much awareness of its conserva- acute, often reflexed on older branchlets and on whip
tion value and an active management programme shoots; stomata on both sides in two marginal lines;
(largely to prevent wildfires) is being implemented. leaf colour dull light green. Pollen cones oblong, 36
Further cutting of trees is prohibited. 1.52mm; microsporophylls peltate, coriaceous,
IUCN: VU (A1c, d) broadly triangular to ovate, bearing 35 abaxial pol-
len sacs near base. Seed cones solitary or grouped
Uses (not clustered) on foliage branchlets, sometimes
terminal on short shoots, maturing in two growing
Willowmore Cedar was used in the past for con- seasons to irregular-subglobose, 1522mm diam.,
struction timber of local farmsteads and for furni- serotinous but eventually wide openening cones.
ture making. Depletion of the resource to a few trees Bract-scale complexes 4 (rarely 6), thick woody,
in inaccessible locations terminated this use and the oblong, max. 25 15 8mm, curved slightly inward;
remaining trees are now protected. In cultivation it outer surface smooth to rugose, not or variably ver-
is only present in a few botanic gardens and in some rucose, with a 14mm long, subapical, upcurved or
private gardens in California and Oregon. It should recurved boss enclosing or exposing the bract apex,
be tried more often in summer-dry regions. (reddish) brown or blackish brown; inner surface
with angular, 34mm long, whitish seed scars at
base, red-brown, blackish towards base. Columella
Widdringtonia whytei Rendle, Trans. Linn. short, thick, angular, to 5mm tall, sometimes dou-
Soc. London, Bot., ser. 2, 4: 60, t. 19. 1894. ble. Seeds 310(18) per cone, ovoid, flattened, 57
Widdringtonia nodiflora (L.) Powrie var. whytei 23mm without wings, blackish brown or black,
(Rendle) Silba, Phytologia 68: 74. 1990. Type: with dull lighter hilum at base; wings 2, of unequal
Malawi: Mulanje District, Mulanje Highlands, Mt. size, up to 3mm wide near seed apex, not joining,
Mulanje, A. Whyte s.n. (holotype BM). Fig. 357 brown, translucent.
Etymology Distribution
This species was named after Alexander Whyte, who S Malawi (Mt. Mulanje).
explored Mt. Mulanje, Malawi, in 1891. TDWG codes: 26 MLW
Ecology the incidence of fires. Illegal timber extraction con-
tinues to be a problem. Another potential threat is the
Widdringtonia whytei is an important to co-dom- invasive pine Pinus patula, which is now being found
inant species in the Afromontane forest on Mt. in both grassland and forest on the mountain. A more
Mulanje, which also includes Podocarpus milanjia- proactive strategy, involving the protection of natural
nus, Cassipourea malosana, Ekebergia capensis, Olea regeneration and perhaps planting, is urgently needed
capensis, Polyscias fulva, Rapanea melanophloeos, to save this species from extinction. Ecologically it
and Xymalos monospora, and in the more fire-prone is at a disadvantage compared with its more adapt-
ecotone (with ericaceous scrub) to grassland, the able congener on Mt. Mulanje, W. nodiflora (Pauw &
closely related species Widdringtonia nodiflora. It Linder, 1997). In past plantings, no distinction seems
is a successional species after fire (periodic fire cli- to have been made between the two species and this 1061
max), but unlike its congener, it does not coppice has to change if future plantings are to successfully
from (fire-caused) stumps and has to regenerate contribute to the restoration of W. whytei.
from seed (Pauw & Linder, 1997). Thickets of Erica IUCN: CR [A4a, ce; B2ab (iv)]
benguelensis which develop after fire offer protec-
tion for cedar seedlings, leading to Widdringtonia Uses
whytei becoming the dominant tree until invading
angiosperms succeed; however these have been pre- The wood of this species is highly valued for construc-
vented from doing so by the next fire at a cycle of tion and building of houses as it is the only sizable
100200 years. Mt. Mulanje is a granitic batholith tree in the family Cupressaceae that occurs naturally
rising through surrounding older sediments. The in a large surrounding region. Its wood, as of other
soils are therefore largely rocky, acidic and shallow members of the family, is decay and insect resistant.
except in colluvial pockets in gorges and valleys. The Large timber yielding trees are becoming increas-
altitudinal range is 18302550m a.s.l. The climate is ingly rare due to overexploitation of this valuable
cool tropical montane, with abundant precipitation, resource for more than a century (Chapman, 1995).
much of it as fog. Attempts at plantation forestry using this species
have been made, but have been much less sucessful
Conservation than the plantations of the exotic species Cupressus
lusitanica which grows faster. It also appears that
This species is acutely threatened with extinction most of these plantations involve a mixture of two
because of excessive felling in the past 100+ years and species (Pauw & Linder, 1997), often with W. whytei
increased fire frequency (Chapman, 1995). Certainly at a disadvantage as it grows slower. The wood of W.
at present only moribund or severely exploited whytei is yellowish brown and of excellent quality for
stands remain, mostly on less accessible sites. The construction, general carpentry and joinery, wood
species is now officially protected on Mt. Mulanje, panelling, flooring, and furniture making. In colo-
but enforcement proves to be difficult as the timber nial times some of its wood found its way to English
fetches high prices (Chapman, 1995; Pauw & Linder, interiors, nowadays there is no export of this timber.
1997). Population increase in villages surrounding the This species does not appear to be in horticultural
mountain massif indirectly affects the remaining pop- use, although it is present in a few botanic gardens
ulations through increased grazing of livestock and in South Africa.
Wollemia W. G. Jones et al., Telopea 6 (23): 173. 1995. Type: Wollemia nobilis
W. G. Jones et al. (Araucariaceae).
Named after the Wollemi National Park (an or obtuse, hypostomatic, deep green above, glaucous
Aboriginal name for the numerous canyons in the below, broad-based and decurrent; repetitive growth
region). units commencing with short scale leaves to 3mm
long, leaves increasing to 28cm long, 25mm wide.
Description Adult upper canopy plagiotropic shoots initially
near-vertical, becoming horizontal and later pendu-
1062 See the species description. lous, leaves opposite or subopposite tetrastichous and
twisted to present adaxial surfaces uppermost, nar-
Distribution rowly oblong, coriaceous, rounded, dull light to mid
green, unequally amphistomatic, broad-based and
As for the species. decurrent; repetitive growth units commencing with
short scale-like leaves to 3mm long, leaves increasing
to 14cm long, 48mm wide. Pollen cones terminal
Wollemia nobilis W. G. Jones et al., Telopea 6 (23): on first-order leafy plagiotropic shoots, to 11cm long,
174. 1995. Type: Australia: New South Wales, 19mm diam., subtended by ca. 8 helically arranged
Wollemi N. P., [exact locality kept secret], W. G. broadly triangular to semicircular bracts 35mm
Jones NSW 362731 (holotype NSW). Fig. 358, 359, long, 35 mm wide. Microsporophylls helically
360 arranged, dark red-brown, peltate, with a raised angu-
lar termination, with 49 elongate, pendulous pollen
Etymology sacs. Seed cones terminal on first-order leafy plagio-
tropic shoots, usually borne on ascending branches
The species epithet nobilis is Latin for noble and above pollen cones, globular to broadly ellipsoidal,
communicates the impression the tree gave its mid green, becoming brown and shedding individual
describers, but also refers obliquely to the discoverer bract-scales at maturity, to 12.5cm long, 10cm diam.
of the first population of the species, David Noble. Bract-scales flattened, laterally winged, 1217mm
long, 1422mm wide, 35mm thick, with a narrowly
Vernacular names triangular apical extension 612mm long, 24mm
wide at base. Seed scales wholly fused with and indis-
Wollemi pine tinguishable from bracts. Seeds circumferentially
winged, pale brown, 711mm long, 57mm wide,
Description 69mm wide including wing, remaining attached to
the scale. Germination epigeal, cotyledons 2.
Evergreen monoecious trees to 40m tall, frequently
coppicing from base; trunks to 1.2m diam, crown Taxonomic notes
slender, columnar, broadest at about a third of total
height. Bark peeling in thin, fragile, equi dimensional The discovery in 1994 of this unknown conifer
dark red-brown scales on younger stems, becoming and the realization that it belonged to the fam-
densely covered with soft and spongy darkening ily Araucariaceae, but did not fit with either of the
nodules or tubercules to 10mm in diam. Orthotropic two genera known in that family, caused consider-
shoots with helically arranged, decurrent, narrowly able excitement among botanists. Further research
triangular leaves 310mm long, 24mm wide at indicated a slightly closer relationship with Agathis
base. Juvenile and lower canopy plagiotropic shoots than with Araucaria (mainly based on molecular
horizontal, leaves opposite or subopposite, distichous data), but many characters are distinct from either
and twisted to present adaxial surfaces uppermost, or similar to one but not the other in a combination
linear to narrowly triangular, chartaceous, rounded unique to this new genus Wollemia. It soon became
apparent that a type of fossil pollen, Dilwynites, a single canyon system, have been found. Extensive
which had long been known to petroleum geologists searches have not yielded any other, more distant
in Australia, belonged to Wollemia and later some stands of this tree, so it is likely that the total popula-
macrofossils were ascribed to it as well, taking the tion does not exceed 100mature trees and is prob-
fossil record back to the Late Cretaceous (Turonian) ably smaller. Counting individuals is difficult as the
as far as is presently known (Kunzmann, 2007). The trees readily coppice from base or root meristems
fossil record for Araucaria is substantially older and several trunks may belong to a single individual.
(middle Jurassic) and extinct representatives of the Access is difficult and the site is still kept a secret,
family Araucariaceae were present in the Triassic. although by now (2016) there must be people who
The chance survival of Wollemia in a gorge sur- have unauthorized knowledge of it. The species
rounded by an environment hostile to it must count occurs entirely within the undeveloped Wollemi 1063
as one of the remarkable events in the history of life. Wilderness of Wollemi National Park. Regeneration
occurs both from seeds and vegetatively and under
Distribution entirely natural conditions the species seems able to
survive. However, effects of human disturbance on
Australia: New South Wales (Wollemi National site and, in the longer term, possible effects of cli-
Park). mate change (e.g. drying of the permanent canyon
TDWG codes: 50 NSW-NS bottom creek) make these small populations, the last
to exist in a long declining trend as evidenced by the
Ecology fossil record, critically endangered.
IUCN: CR [B1ab (iii), B2ab (iii)]
Emergent trees above warm temperate rainfor-
est dominated by Ceratopetalum apetalum and Uses
Doryphora sassafras, in a deep sheltered gorge
surrounded by sandstone cliffs of the Triassic An extensive propagation and cultivation pro-
Narrabeen Group. Tall eucalypt woodland domi- gramme as well as marketing and PR have been
nated by Eucalyptus piperita is adjacent on can- launched shortly after the discovery of this species.
yon walls and steep slopes, giving way to dry open It has been available for sale since 2005 as a living
woodland up-slope. The soil is sandstone-derived fossil in other words a curiosity; however, it appears
boulder alluvium, with high organic matter, some to be suitable for large gardens in milder climates
shale component, and a substantial basalt wash from (tolerating occasional light frosts, perhaps to zone 8
higher reaches of small tributary canyons. The local in Europe) and is an easy grower readily taking root
microclimate is wet, with a permanent creek and an from cuttings or scions. It is now well established in
understorey dominated by ferns. horticulture and available from nurseries in many
countries.
Conservation
Plate 42. Xanthocyparis vietnamensis. 1. Branch with adult foliage and seed cones. 2. Branch with
juvenile and adult foliage. 3. Adult leaves. 4. Juvenile leaves. 5. Surfaces of juvenile leaf. 6. Pollen cone.
7.Microsporophyll with pollen sacs. 8. Seed cones. 9. Seeds.
Description with whitish hilum at base; wings 0.51mm wide,
thin menbranous, lighter coloured.
Trees to 1015m tall; trunk monopodial, up to 50cm
d.b.h. Bark on the trunk of larger trees becoming Distribution
soft and fibrous, brown to grey-brown, exfoliating
in numerous thin strips. Branches spreading more Viet Nam: N Viet Nam, Ha Giang Province, very
or less horizontally; forming a pyramidal crown in localized in the Bat Dai Son mountain system near
young trees but a spreading, irregular or flat-topped the Chinese border.
crown in old trees. Foliage in crowns of both small TDWG codes: 41 VIE
and larger trees of two or three types: predominantly
1068 with adult leaves, also with juvenile leaves, often Ecology
also with transitional leaves. Foliage sprays with
adult leaves flattened; ultimate branchlets spreading This species occurs in mixed angiosperm-conifer
at 3045, 520mm long, 1.53mm wide. Foliage forest with the conifers Amentotaxus argotaenia,
sprays with juvenile leaves bushy, less branched; Nageia wallichiana, Pseudotsuga sinensis (domi-
branchlets with transitional leaves more like those nant), Podocarpus brevifolius, and Taxus chinensis
with adult leaves. Adult leaves decussate, short and numerous, mostly small leaved angiosperms.
decurrent, imbricate, dimorphic, on (sub)ultimate The conifers appear to be restricted to the ridges and
branchlets 1.53 11.3mm (the laterals slightly lon- summits of the mountains. Dominant among angio-
ger than the facials); facials narrowly ovate-rhombic, sperms are species of Acer, Carpinus, Lithocarpus,
keeled, more or less appressed; laterals conduplicate, Quercus, and Ulmus; frequent are Pistacea wein-
spreading free from the leaf above at ca. 30, straight mannifolia and Platycarya strobilacea. In a second
or falcate; margins of both types minutely denticu- stratum under the ca. 20m tall canopy occur fre-
late except towards acute or pungent apex; stomata quently species of Elaeocarpus, Eriobotrya, Sorbus,
on adult leaves inconspicuous, mostly adaxial; Schefflera, and many others. Shrubs and herbs
glands inconspicuous. Transitional leaves similar to abound, among the latter are numerous species of
adult leaves but longer (57mm), lanceolate; laterals Orchidaceae, terrestrial as well as epiphytic, some-
spreading wider at 45. Juvenile leaves in whorls of times determining the aspect of the ground cover
four, decurrent, with distal part spreading at nearly vegetation (Averyanov et al., 2002). Ferns and espe-
90 and proximal decurrent part 45mm long; cially bryophytes are similarly abundant, both as
distal free parts 1520 1,52mm, linear; margins lithophytes and as epiphytes. The limestone ridges
entire, tapering to a fine point; stomata in two whit- on which Xanthocyparis occurs are extremely
ish bands on abaxial side only. Pollen cones 2.54 eroded, composed of hard, marble-like rock out-
22.5mm, oval-terete; microsporophylls 1012, crops interspersed with thin soil pockets. The alti-
peltate, ca. 1 1mm, each bearing abaxially 2(3) tude of the ridges where this conifer grows is from
yellow pollen sacs. Seed cones green, turning dark 1050m to 1330m a.s.l. The climate is subtropical, but
or dull brown, subglobose, 911 1012mm when damp and wet much of the year (cloud forest).
open, some more or less persistent after seed disper-
sal. Bract-scale complexes 4(6), decussate, fused Conservation
at base; upper and larger pair connate distally, then
spreading, valvate to subpeltate; lower pair oblong; This species is restricted to a few localities in close
all widest distally, with rounded but irregular upper proximity, mostly in inaccessible sites on steep lime-
margin; outer surface rugose or radiately furrowed stone ridges. In some localities trees are cut, pre-
from a prominent, 12.5mm long, subapical umbo; sumably for local use, in others no such disturbance
inner surface reddish brown marked proximally could be found, possibly due to inaccessibility. The
with whitish seed scars; a small columella present total extent of occurrence (EOO) appears to be less
or absent in the centre. Seeds up to 89 per cone, than 50 km2 within which the species occupies ridge
ovoid or irregular, flattened, 4.56 45mm includ- tops and summits only. Although difficult to assess
ing two lateral wings, light brown or reddish brown, precisely due to the difficult terrain, the total area
of occupancy (AOO) is estimated to be less than Uses
10 km2 (Averyanov et al., 2002). Regeneration has
been observed in habitat. An earlier estimate (Farjon The Golden cypress produces fine, yellowish brown,
et al., 2002) put this species at Critically Endangered very hard, fragrant timber. The majority of trees seen
(CR), as did Averyanov et al. (2002), but objective on the ridges are small or contorted and would not
evidence of serious decline has not been given. be suitable for timber of any trade value other than
IUCN: EN [B1ab (iii), B2ab (iii)] local use. To date, no direct information of its actual
or potential uses is available. Cultivation is being
undertaken locally with an objective of ex situ con-
servation; a few botanic gardens in the UK and USA
also grow it for this reason. 1069
APPENDIX
In this appendix to the second edition of A Hand leaves on juvenile and adult plants; midrib nar-
book of the Worlds Conifers newly described and row (<0.5mm wide) not or only slightly raised on
accepted species are presented that were published both surfaces, not forming a groove on the adax-
between the appearances of the two editions. To pres- ial (upper) surface; apex more or less obtuse or
ent them in this place in the book avoids disruptive sub-acute, base acuneate. Pollen cones solitary or 1071
and complicated changes that would have to be made paired, grouped but not densely clustered with up
in an alphabetically arranged account of the species to 3close together, pedicellate, the pedicel of indi-
were they to be inserted in their appropriate place. vidual cones 39mm long; pollen cones 2036 x
23.5mm, yellow(ish), narrowly cylindrical, curved,
Podocarpaceae with numerous microsporophylls with a deltate free
part. Seed cones paired, lateral on a reproductive
shoot on long peduncles (to 15mm) longer than
Podocarpus orarius R. R. Mill & M. Whiting, Gard. the receptacle; foliola 2.5mm long, soon caducous;
Bull. Singapore 6 4 (1): 176. 2012. Type: Solomon receptacle asymmetrical, rectangular-ellipsoid when
Islands, Choiseul, Loloko District, mainland opp. ripe, 1013.5mm long, 8.510.5mm wide, turning
Bembalama Island, F. Pitisopa et al. 7 (holotype E). yellow then deep red or vermillion at maturity. Seed
subglobose, 1112 x 99.5mm, laterally compressed,
Podocarpus spathoides de Laub. var. solomonensis not crested; seed coat and epimatium olive green,
Silba, J. Int. Conifer Preserv. Soc. 7 (1): 39. 2000. rugose when dry.
Etymology Distribution
The epithet is Latin for coastal and refers to the Solomon Islands, Vanuatu (Erromango).
most common habitat observed by collectors of TDWG codes: 43 SOL-SO 60 VAN
specimens.
Ecology
Vernacular names
Mostly occurring on steep slopes in coastal rain-
Dengali seems to be the name consistently applied forest (primary or also secondary) often associated
to this species. with species of Gymnostoma (Casuarinaceae), from
near sea level to around 450m. Possibly also present
Description in interior lower montane rainforest.
abaxial In a position removed from an axis or the seeds are consequently enclosed in an ovary
stem (also dorsal). which develops into a fruit of some kind (com-
acicular Needle-shaped; long, narrow, straight or pare gymnosperms).
slightly curved, slender, and not or only slightly anthesis (in gymnosperms) The action or period
flattened. of pollen dispersal and pollen reception 1073
acroscopic Facing or directed towards the apex (flowering).
(here of the foliage branch). AOO (area of occupancy) Term used in IUCN
acuminate Tapering gradually to a well defined, Red List criteria to describe the estimated area
sharp point. (in hectares or square kilometres) actually cov-
acute Sharply pointed. ered or occupied by a species. Compare EOO.
adaxial In a position nearest an axis or stem. apex In plant morphology, the terminal point of
adnate Joined with another organ of a different a growing structure like a shoot or a leaf. The
kind; e.g. the seed wing with a seed (compare adjective is apical; a conifer cone that is apical
connate). grows at the apex of a shoot (but very few are).
adult leaves Here the type of green leaves that apiculate Ending abruptly in a small, distinct
appears and is predominant on primary (or point.
old) branches of mature trees, usually in the apiculus A small, abrupt and/or distinct point at
crown and fully exposed to sunlight (compare the apex.
juvenile leaves). apophysis The well defined apical part of ovulif-
afforestation The planting or seeding of trees in an erous scales in the seed cone marking the first
attempt to create forest. What is meant with the stage of growth in Pinus; it is rarely differenti-
term forest has changed over time; in this con- ated in the other genera of Pinaceae.
text it now usually refers to plantations used as appressed Pressed close to or lying flat against
a source of timber. another organ (e.g. the shoot or other scales).
aggregate Grouped closely or densely together, arboretum (plural: arboreta) A park or garden
clustered. dedicated to a living collection of planted trees
allopatric The occurrence of populations or spe- and shrubs.
cies in geographically separated areas. archegonia (singular archegonium) In gymno-
amenity planting Planting of ornamentals and sperms, the unfertilized female sex organs, i.e.
trees for general use in public spaces, e.g. street the parts of the ovule that contain the female
trees, flower borders, and shrubs in parks, high- nuclei with a haploid number of chromosomes.
way intersection plantings, etc. Gymnosperms have several archegonia per
amphistomatic Stomata on both abaxial and ovule.
adaxial sides, on equifacial leaves on all three aril(Latin arillus) A fleshy or succulent (cup-like)
or four faces; equally distributed, or more often structure that completely or partially envelops a
with two primary bands on two adjacent faces seed and is derived from the integument (com-
(compare epistomatic and hypostomatic). pare epimatium and drupe).
ampulliform Shaped like a flask (Latin ampulla), ascending Rising obliquely or curving upward.
with a wide body and a narrowing neck below a.s.l. Above sea level.
the apex. asperous Harsh to touch or handle; here due to
anastomosing Conneting so as to form a network rigidity of pungent leaves.
(compare reticulate). association (in vegetation science) A fundamental
angiosperms Flowering seed plants with ovules unit of vegetation, which can be distinguished
attached to carpels, developing into seeds sur- from other such units on the grounds of its flo-
rounded by a pericarp formed by the carpels; ristic composition.
assurgent Bent near the base, then steeply rising to boreal(Greek Boreas = god of the north wind) Of
a (nearly) vertical position. northern regions of the world.
auriculate (auricled) With ear-like lobes at the bract A leaf-like or scale-like appendage, usually
base or lower sides. subtending a fertile structure such as a seed
axil The angle between a bract, branch, or leaf and scale. In some conifers, the seed scale is fused
the axis from which it arises (i.e. rachis, shoot, with the bract, rudimentary, or altogether
branch, or stem). absent, by which the ovules (seeds) are situated
axillary Positioned in the axil. in the axil of bract and cone axis, or sometimes
basionym The name-bringing or epithet-bringing solitary on the apex of the axis.
synonym of a newly proposed combination. bract-scale complex Term used in Cupressaceae
1074 bifacial With two distinct, opposite faces. If flat- for the scales of seed cones; these are technically
tened in that way, the organ, e.g. a leaf, has its bracts, but transformed by intercalary growth
margins bordering these faces (see also dorsi- to form the cone scales and enclose the seeds.
ventral and compare bilateral). In some complexes there is a rudimentary seed
bifid Deeply cleft at apex, so as to form two sharp scale present (in related Sciadopityaceae it is
teeth. fully developed), in most the true seed scale
bijugate An arrangement (of leaves) in which does not develop.
there is a doubling of the numbers 2 + 3 in nor- buttressed Shaped like a buttress; here the broad-
mal (unijugate) spiral arrangements; the biju- ened base of a tree trunk, usually continued
gate arrangement of 4 + 6 parastichies is in downward in the above-ground parts of roots.
fact an opposite-decussate arrangement with a caducous Falling off at an early stage.
twist. callous Being hardened and thickened.
bilateral A symmetry in which similar parts (sides) cambium Primary tissue (see meristem) forming
are arranged on opposite sides of a median axis; secondary tissue. There are two cambia: one in
in leaves: flattened with two equal sides (not the outer part of the stem (cork cambium) and
faces; compare bifacial and dorsiventral). one just outside the wood of the stem; it forms
bilobate With two lobes; usually a terminal part secondary wood on its inner side and second-
divided in two. ary bark on its outer side.
binomial A name of a thing consisting of two sep- canaliculate With a single, longitudinal groove.
arate words; in biology it refers to the names of candelabra crown A tree crown conforming to
species, which are always the name of a genus Rauhs model but with few first order branches
written with a capital first letter followed by an in pseudo-whorls restricted to the upper part of
epithet written in lower case. the ited to the ends of these.
biodiversity The total number of taxa (or some- canopy The upper crown level in a closed forest,
times genetic diversity) known to exist in a where the crowns of the tallest trees meet to
given area; the greater this figure turns out to form a more or less continuous cover.
be, the higher the regions biodiversity. capitula (Latin = small head; plural: capitulae) A
biogeography The study or science of the geo- dense, compact cluster of flowers. In conifers,
graphical distribution of organisms and its this refers to pollen cones, not flowers.
causes and history. carinate Keeled; ridged lengthwise.
biome A major ecological community type, related cataphyll A reduced, scaly primary leaf; in pines
a to major climatic zone. they form winter buds, enclosing an entire shoot
biota (Latinised form of Greek biot = life) The (with leaves and internodes not yet elongated)
totality of life (plants, animals, fungi etc.) in a of the next growing season. On expansion of
region. the shoot each cataphyll subtends a dwarf shoot
bisaccate(in pollen) Having two sacci or air blad- with secondary leaves (needles).
ders giving the corpus of the pollen buoyancy caudate Elongated, resembling a tail, or tail-like
in the air. appendage.
bole The straight trunk of a tree.
chaparral Term used in the southwest of the USA variables) of at least three different organisms
and Mexico to describe a dense vegetation of (representing species or higher ranks of taxa),
shrubs, some of which are evergreen, which showing the inferred phylogenetic relationships
grows in coastal regions with a dry summer and of these characters based on cladistic principles.
a moist winter climate, and often forms a dis- clavate Club-shaped, i.e. with a broader or thicker
tinct vegetation belt (see also maquis). distal part.
character A feature of an organism which has climax A supposedly terminal stage in the succes-
variable properties. sion of vegetation types from pioneer coloniz-
character state The variable property of a character. ers to mature vegetation. In fact, all stages in
Leaf length is a character, the actual measure- the succession are temporary since they are all
ment given in cm or mm is the character state. subjected to natural disturbances, including the 1075
chlorophyll A green pigment, contained in chlo- terminal or climax stage.
roplasts in leaf or sometimes stem tissue of cline A graded series of morphological (or physi-
plants, responsible for photosynthesis. ological) differences exhibited by a group of
chloroplast A microscopic body (organelle) inside related organisms, usually along a line of envi-
a plant cell that contains chlorophyll. ronmental or geographic transition.
chromosome A usually linear body of chromatids, codominant Dominating together with other spe-
containing nucleoprotein and DNA, the mol- cies in a given vegetation; trees are codominant
ecule responsible for the hereditary code, in the when they form part of the main canopy of a
cell nucleus; the number of chromosomes usu- forest.
ally remains constant in a species. columella (Latin = little column) In conifers a
ciliate Fringed with regularly arranged hairs on spiky protrusion in the centre (axis) of the
the margin; the hairs are usually fine and of inside of a seed cone, especially prominent in
equal length. those of the genus Callitris (Cupressaceae).
CITES Convention on the International Trade in concave Hollowed or rounded inward like the
Endangered Species of Wild Fauna and Flora. inside of a bowl (compare convex).
The signatories to this convention (1976) are conduplicate Folded together lengthwise.
nation states; they decide at regular conferences cone In conifers, referring to a structure bearing
of the parties to restrict or prohibit cross-bor- either pollen sacs containing pollen, or ovules,
der trade (= movement by people) of species which after fertilization may develop into seeds.
listed on Appendices I, II, or III (with I the most Conifer cones can be simple, consisting of an
restrictive). axis with one type of scales, or compound, with
clade A section of a cladogram which indicates a two types of scales, only one of which bears the
monophyletic (holophyletic) group of termi- ovules. Pollen cones in conifers are simple; seed
nal taxa (2). cones can be compound or simple.
cladistics A formalised method of character com- congener A species in the same genus as another
parison developed by the German entomologist species.
Willi Hennig in 1950; it only considers shared conifers (cone-bearers) To this taxonomic group
derived characters to inform phylogeny and its belong those woody gymnospermous plants
aim is to construct a hypothesis of phylogenetic (usually trees) of which the seed bearing scales
relationships between groups of organisms are united in a cone. The conifers (including
(usually recognized taxa). forms with extremely reduced cones, such as
cladode A branch-like or shoot-like organ not Podocarpus, or those with no apparent cones,
being a green leaf, but functioning as such such as Taxus) are a monophyletic group of
(compare phylloclade). In Sciadopitys true plants derived from a single common ancestor.
leaves form small scales in the axil of which connate Joined with another organ of a simi-
arise needle-like, seemingly paired, and fused lar kind, e.g. a leaf with another leaf (compare
cladodes, functioning as leaves. adnate).
cladogram A branching diagram resulting from convergence The evolution towards similar mor-
a cladistic analysis of characters (i.e. discrete phology not as a result of common descent, but
of adaptation to similar environments or life and projecting-lipped drinking vessel sup-
strategies. ported on a standard.
convex Curved or rounded like the exterior of a d.b.h. Diameter at breast height; used to measure
sphere or circle; here used for the shape of the the girth of a tree trunk above a buttressed base
cone scales as viewed from the abaxial side or roots; for very large buttressed trees mea-
(compare concave). surement should be taken at actually higher
copal A kind of resin; commercially referring to (than the usual ca. 1.3m) levels.
resins of leguminous and araucarian plants that deciduous Falling off or shed seasonally or at a
produce hard varnishes. certain stage of development in the life cycle.
coppice Regrowth from the stump of a tree after decumbent (Latin = lie down) Growing low over
1076 cutting (or burning), producing new stems and the ground or surface of a rock.
branches. The stump after cutting is called cop- decurrent Running down a stem from a leaf
pice stool; in many species the stems will regrow base.
even after repeated cutting. decurved Curved downward (compare recurved).
cordate Heart-shaped (subcordate = somewhat decussate Occurring in alternating pairs; leaves
heart-shaped). are decussate if one opposite pair is placed
coriaceous(Latin corium = skin) Resembling slightly higher than the adjacent opposite pair
leather: leaves with a leathery, tough texture, and oriented 90 from it (compare ternate).
mainly from thickened dermal layers. deflected(Latin deflexus) Bent or turned abruptly
cortex Layer of` cells (parenchyma) in plant roots downwards.
and stems, situated between the central vascular dehiscent Referring to the manner of opening of
cylinder and the epidermis or surface layer. a vessel containing seeds or spores; in conifers
costate Ribbed, having one or more primary veins it pertains to the pollen sacs or to some types of
or ridges. globose seed cone.
cotyledons First, or embryo leaves, as found deltoid triangular with the sides more or less equal
in the seed and which (usually) unfold after and the apex uppermost.
germination. dendrochronology The science of dating events
crenate (crenately lobed) Scalloped, lobed like the and variations in the environment in former
surface of a scallop. periods by comparative study of growth rings in
cultivar A form of a plant (not a botanical vari- the woody stems of trees.
ety!) in cultivation, the characteristics of which dendrological Of dendrology, the science or study
are perpetuated by vegetative propagation. of trees.
cuneate Wedge-shaped; in the descriptions often dentate Having the margin cut with sharp, salient
combinations are used, e.g. obovate-cune- teeth, directed outward or straight.
ate, to describe either a range of shapes or denticulate Finely or minutely dentate.
intermediates. dermal Of the skin (Greek derma = skin), in coni-
cupular Cup-like, similar in shape to the cupule of fers it can refer to bark or more commonly to
an acorn. leaf surfaces or outer layers (see epidermis).
cusp The usually well defined pointed apical determinate Referring to growth of a structure
extension of a leaf (but here also of the bracts which definitely ends with the completion of
of seed cones). that structure. Conifer cones represent shoots
cuspidate With a cusp, i.e. a (short or long) well with determinate growth, while foliage shoots
defined and tapering apical point. can and often will resume growth in the next
cuticular Referring to a usually waxy surface layer growing season. In some abnormal cases, cones
on the exterior of the epidermis, consisting of a are not determinate and a foliage shoot may
protective substance (cutine); it causes the shine grow from their apex (compare proliferation).
on conifer leaves. dichotomous Like a system of branching in which
cyathiform Shaped like a beaker (in outline) the main axis forks repeatedly into two branches
(Latin cyathus = beaker), i.e. a wide-mouthed (dichotomy).
dilated(Latin dilatatus) Expanded, broadened, or ecosystem The supposed totality of the functional
widened (sometimes also swollen). relationships of organisms with their environ-
dimorphic Of two forms or shapes; an organ, e.g. a ment and with each other living in a given area
leaf, has two different forms (2character states) and habitat.
occurring on the same plant or shoot (compare ecotone The transitional zone between two types
monomorphic and polymorphic). of vegetation or ecological communities, usu-
dioeciousIn gymnosperms this pertains to ally with (competing) species of both types and
female and male reproductive organs, com- often with species not occurring in either of the
monly cones, being present on different plants two types.
(see also monoecious). ecotype A plant form with morphological char-
diploid Cells containing two sets of chromosomes acters that are modified by the environment in 1077
(2n = x). The nuclei of these cells contain sets which the individual plant grows; such char-
of paired chromosomes of equal form and size. acters are not inherited and therefore are not a
Such cells have a doubling of the base (haploid) good basis for taxonomic distinction.
number of chromosomes, the result of sexual emarginate Shallowly notched at the apex.
fusion of female and male haploid nuclei. A rare embryo (in plants) The young sporophyte of a
further doubling of the diploid number leads to seed plant, usually consisting of a very small
tetraploidy (Greek tetra = four) and in one spe- plant with a primary bud (plumule), radicle,
cies (Sequoia sempervirens) hexaploids (Greek and cotyledons.
hexa = six) were found, which amounts to a tri- emend.(Latin emendavit = changed by) Indicating
pling of the diploid number. an alteration of content or circumscription, fol-
disjunct (of distribution) Occurring in areas lowed by citation of the author(s) responsible
widely separated from each other. for the change.
dispersal The spreading of organisms to areas emergent In the context of trees referring to indi-
other than where they originated. viduals rising above the forest canopy.
distal Occurring nearest the apex or terminal part endemic (endemism) Referring to the occurrence
(compare proximal). of a taxon in a limited area for example, on
distichous Disposed in two vertical or upright rows. a single island. Taxa limited to a single coun-
divaricate (from Latin divaricare = to straddle try are also considered endemic, although the
as in riding a horse) Divergent branching, with term becomes fairly meaningless biologically
two branches splitting from the apex of a single for widespread species limited to countries that
branch or stem. cover nearly half a continent.
DNA (deoxyribo nucleic acid) The macro-mol- endophyte An organism that grows within (in the
ecule in cells of organisms that contains the context of trees a branch, root, or stem).
genetic information and acts as a template to entire Without teething or division, with even
build new cells and therby (multicellular) new margin.
organisms (compare RNA). EOO (extent of occurrence). Term used in IUCN
dolabriform Shaped like an axe. Red List criteria to estimate the total range
dorsiventral The condition of possessing upper (inkm2) within which a species is found to
and lower faces of an organ. If flattened in that occur naturally (excluding introductions), usu-
way, the organ, e.g. a leaf, has its margins bor- ally by connecting the outermost points of pres-
dering these faces (see also bifacial). ent occurrence. Within the EOO is the AOO,
drupe A fleshy, one-seeded, indehiscent fruit which is always smaller.
with a stony endocarp (e.g. cherry, peach). It is epicormic (Sprouting) from dormant buds on the
derived from the carpels of angiosperms (com- trunk lying within the bark.
pare aril and epimatium). epidermis The true cellular outer skin of a plant
ecology The study or science of ecosystems, or of (leaf) below the cuticle, made up of a single
the functional relationships of organisms with layer of cells.
their environment and with each other.
epigeal germinationThe hypocotyl of the seed- of the genetic makeup of individuals in popu-
ling elongates and lifts the cotyledons above lations, primarily through genetic mutation,
the soil surface, where they unfold (compare separation (isolation) and natural selection,
hypogeal). resulting in new species. The overall tendency is
epimatium The outer layer of tissue that (partly) divergence, interrupted by extinction, causing
covers a seed in many species of the family major change over geological time.
Podocarpaceae; it is derived from the seed scale exfoliating Here: the dead outer layers of bark
(compare aril and drupe). coming off in scales, sheets, or papery flakes.
epiphyte A plant that grows upon another plant exserted Protruding; here usually pertaining to
(usually a tree) using it for support only. Plants the bracts of seed cones projecting beyond the
1078 that are accidentally growing on trees because upper margins of the seed scales.
seed germinated there, but of which the usual ex situ (Latin = outside [its] place) In the con-
habitat is terrestrial, are not (obligate) epiphytes text of plant species conservation, any effort to
in an ecological sense. No obligate coniferous conserve individuals by growing them in locali-
epiphytes are known. ties outside the natural range of the species is
episodal Occurring at irregular intervals in time considered ex situ conservation (compare in
(episodes). Here referring to stochastic distur- situ).
bance events from natural causes that can reset exuding Issuing a substance, such as resin,
the successional phases of forest regeneration in through a surface layer of tissue; the product is
an area to a pioneer phase. called exudate.
epistomatic Stomata (almost) restricted to the facials (facial leaves) The paired, opposite scale
adaxial side of the leaves (the dorsal side mor- leaves on a plagiotropic foliage branchlet (esp.
phologically); on plagiotropic lateral shoots the in the family Cupressaceae) that face either up
stomata are facing downward by orientation or down, they are flanked by the differently
of the leaf bases, petioles or pulvini (compare shaped laterals.
hypostomatic). falcate Sickle-shaped (falcate-linear = with two
epitype(Latin epitypus) A specimen or illustra- nearly parallel sides).
tion selected in addition to a holotype (or neo- fastigiate A branching habit in trees in which the
type or lectotype) to serve as an interpretative branches are directed upwards (ascending), not
type if the designated type material or other spreading out from the trunk.
original material cannot be identified for the farinose Covered with a flowery or meal-like
purpose of precise application of the name of powder.
a taxon. ferruginous With the reddish brown colour of
equifacial With equal faces; i.e. the 3 or 4 faces of rusty iron.
those leaves that are quadrangular or rhombic figured In sawn and planed or turned and pol-
in cross-section. ished wood the decorative patterns and shapes
ericoid (as in Erica) Belonging to (or similar to) caused by growth rings and other structures in
Ericaceae; also an adjective used to describe relation to the plane of sawing or turning.
a vegetation dominated by such plants, or by fimbriate Fringed with long hairs.
those with a similar life form (i.e. evergreen flabellate Fan-shaped (though in cone scales
winter hardy shrubs). of conifers not entirely flat but more or less
erose Irregularly toothed, as if gnawed; here often concavo-convex).
used in conjunction with denticulate. flaccid Weak and lax, lacking firmness.
eutrophic Adjective referring to an environment flexuousLatin flexuosus) Bent alternately in oppo-
(usually a body of water) rich in dissolved nutri- site directions, in zigzag fashion.
ents available to plants. flushing (of leaves) The more or less abrupt
evolution Gradual change (as contrasted with rev- unfolding from winter buds or just growth of
olution). In biology the term is mostly used in a new foliage leaves. In tropical trees, these young
Darwinian sense and refers to (gradual) change leaves are often coloured pink or red, or a
ifferent green from leaves that have been
d graft The artificial growing connection of two
exposed longer. parts of different plants, involving a root stock
foliate Leaf-like, i.e. shaped like a leaf as in a book and the grafted cutting or scion with foliage.
(Latin folio). Leaves of plants (conifers) can have The resulting plant minus the rootstock is also
shapes different from foliate, such as acicular. referred to as a graft.
foliola (Latin sing. foliolum) Little leaves or leaf- Greggs Paradox This paradox seems to violate the
lets. A term used to describe two bract-like hierarchical structure of biological classifica-
appendages subtending the receptacle in seed tion: Ginkgo = genus Ginkgo biloba = species
cones of the genus Podocarpus. Ginkgo = Ginkgo biloba genus = species.
frondose Resembling a pinnately compound fern The way out of this paradox is to postulate
leaf (frond). extinct species and only one surviving to the 1079
fusiform Spindle-shaped; tapering at both ends. present. For Ginkgo there is abundant evidence
fynbos A vegetation formation occurring in the of other species in the fossil record, for other
Cape Province of South Africa, characterized by genera with one extant species it is possible that
a low, open structure and the abundance of eri- these will be found in future. [From J. R. Gregg
coid, proteoid, restioid and geophyte life forms (1954). The Language of Taxonomy. Columbia
on nutrient-poor soils. University Press, New York.]
gametophyte The generation (in spore-repro- gymnosperms Non-flowering seed plants with
ducing plants) that bears the sex organs; in ovules directly exposed to air (naked); the
gymnosperms strongly reduced to the micro- seeds may be exposed also but are usually
gametophyte in the pollen grain and the mega- enclosed within a cone or by an aril or epima-
gametophyte in an ovule or seed; the latter tium; neither is technically a fruit (compare
refers to the tissue that embeds the (female) angiosperms).
archegonia within an ovule; after fertilization of habit(Latin habitus = condition, character)
the archegonia the ovule develops into a seed. Characteristic mode of growth or occurrence.
The megagametophyte then has the same func- habitat The place or type of site where a plant (or
tion as the endosperm in seeds of angiosperms, animal) naturally or normally grows and lives
but it has a different origin (homology). and to the natural conditions of which it is
gene A bit of DNA (sequence of nucleotides) cod- adapted.
ing for a protein, or part of a protein. hardpan A compact soil surface inhibiting perco-
genome The total of genetic or hereditary infor- lation of rain water to deeper levels where tree
mation present in the DNA of an organism in roots could take it up. Such soils are common in
the form of the sequence of its nucleotides. (semi) arid regions.
genotype The genetic constitution of an individual heartwood The inner section of the wood of a tree,
or a group of organisms. with dead wood cells through which no water is
GIS A geographic information system (GIS) transported.
integrates hardware, software, and data for
helical(Latin helix) Arranged or formed in an
capturing, managing, analyzing, and display- upward spiral (see also spiral).
ing all forms of geographically referenced heterosis A marked capacity for rapid growth
information. often shown by cross-bred animals or plants
glabrescentBecoming glabrous with age. (hybrid vigour).
glabrous Devoid of hairs of any kind. heterotypic (synonym) A synonym based on a dif-
glandular Functioning as a gland, i.e. secreting a ferent type. Such synonymy results from the
biochemical substance. merging of different taxa into one by taxono-
glaucous Covered with a blue-green or blue-white mists (compare homotypic).
bloom (see also pruinose). hexagonal Shaped like a hexagon, i.e. a polygon
globose Nearly spherical (compare globular). with six angles and six sides.
globular Spherical, round (compare globose). hilum A mark on a seed left by its connection to
glutinous Sticky, with the quality of glue. the tissue from which it developed.
hirsute Covered with coarse or shaggy hairs. dermis; it is discontinuous at the stomata and
holotype(Latin holotypus) The one specimen (or otherwise continuous, intermittent or absent.
illustration) used or designated by an author hypogeal germinationThe hypocotyl of the seed-
as the nomenclatural type of a species or lesser ling does not elongate, the cotyledons remain
taxon at the time of establishing it. infolded below the soil surface and only the true
homologous Referring to shared characters in two stem of the seedling emerges (compare epigeal).
or more species due to common descent, or to hypostomatic Stomata (almost) restricted to the
modifications evolved from such commonly abaxial side of the leaves, with two primary
inherited characters. bands separated by a midrib in most species; on
homonym A name of an organism spelled exactly plagiotropic lateral shoots the stomata are fac-
1080 the same as the name of another organism. ing downward by a twisting of the leaf bases or
Homonyms within the jurisdictions of the sev- petioles (compare epistomatic).
eral codes of biological nomenclature are not ICBN International Code of Botanical Nomen
allowed and the rule of priority dictates that clature. A set of internationally accepted rules
the name shall be applied to the earliest named and regulations governing the naming of plant
plant or fungus (in case of the Botanical Code) (and fungal) taxa, decided at International
and that the other plant or fungus must be Botanical Congresses held at six year intervals
renamed. and organised by the International Association
homoplasy (homoplasious) A term for a character for Plant Taxonomy (IAPT), which organisation
that is not a shared derived character (synapo- publishes these rules in the Botanical Code (at
morphy) in a phylogenetic sense. Homoplasious the time of publication of this Handbook the
characters (or character states) appear in more Vienna Code of 2006). Its rulings are binding
than one clade of an inferred phylogenetic tree in plant taxonomy.
(cladogram) and their similarity may result igneous Referring to rock that was formed from
from convergent or parallel evolution, rather molten material (magma) in the interior of the
than from common descent with modification earth; it may have solidified below as well as on
or transformation. the surface.
homotypic (synonym) A synonym based on the imbricate Closely overlapping, as thatches or tiles
same type. Such synonymy results either from on a roof (compare tesselate and valvate).
the strict application of the Botanical Code incertae sedis (Latin = place uncertain) A heading
(ICBN) or from a change in rank of a particular under which taxonomists place names of taxa
taxon. Only the latter procedure involves taxo- for which the original description is unclear in
nomic decision making (compare heterotypic). the context of present knowledge and of which
hort.(Latin hortulanorum) Of gardeners (hor- no good original material exists to determine
ticulturally); here added to a name used in a their identity.
(botanic) garden, but not formally published. If indehiscent Generally: a fruit that does not open.
followed by ex and a name and reference, its In conifer seed cones: the scales remain con-
subsequent formal publication is given. nate, so the cones will not open to release the
hybrid A product of propagation between two seeds.
individuals belonging to different taxonomic indigenous Naturally occurring in an area or a
units; hybridization can be naturally or arti- country, without having been transported there
ficially induced. Some species of conifers are (intentionally or unintentionally) by people.
believed to be of hybrid origin; their botanical inflorescence A branching system supporting
names are marked with a sign. flowers but usually not leaves.
hypocotyl Section of a seedling between radix in litt.(Latin: in litteris) In a letter or written note.
(primary root) and cotyledons, developing into Citations given with this adjective are written
the stem. but unpublished sources.
hypodermis Layer of one or more rows of thick in sicco (Latin = in a dried state) Preserved as a
walled cells, situated directly beneath the epi- herbarium specimen, of which the character
states may markedly differ from living or fresh sometimes on reiterations; these leaves are dis-
material. tinct from adult leaves, but intermediate forms
in situ (Latin = inside [its] place) In the context may occur.
of plant species conservation, any effort to con- karst A deeply eroded formation of marine lime-
serve individuals by protecting them or their stone, of which surfaces are partly dissolved by
habitat within the natural range of the species rain water containing CO2 and where acidic
is considered in situ conservation (compare ex groundwater dissolves caves; drainage in karst
situ). landscapes is largely underground. In tropical
integument Enveloping layer of the seed bud or climates karst landscapes develop tower hills or
ovule, which is open on one side (micropyle) mountains.
and, after fertilization, develops into the seed kerangas Heath forest, i.e. a more or less stunted 1081
coat. (but sometimes tall), open type of forest grow-
intercalary growth Growth inserted between ing on nutrient-poor, sandy (often nearly white)
other parts; in the bract-scale complexes of the soil in tropical lowlands of Malesia.
seed cones of Cupressaceae this occurs inside the keystone species A species in an ecosystem upon
bracts (i.e. between the dermal layers) enlarging which many species depend that characterize
them in various ways to form the cone scales. that ecosystem and make it function. Removal
internal (as of resin canals or ducts) Inwardly; of the keystone species destroys that particular
here a position of the resin canals in the leaves ecosystem.
bordering the endodermis. knot In sawn wood the remainder of a branch in
introgression The entry or introduction of genes the stem of a tree, visible as a darker, rounded
from one gene complex (e.g. a species) into bit of wood with a different grain.
another. K-strategy Also known as K-selection; in plant
invers-dorsal A position of the leaves whereby ecology, this refers to trees with relatively slow
the abaxial (dorsal) side is inversed by twist- growth rates but persistence (longevity) in later
ing of the leaf basis or petiole to face downward phases of vegetation succession. Its opposite,
(below). r-selection applies to trees that colonize early
inverted A position turned about 90 from an ear- phases (pioneer species), grow rapidly but are
lier position; inverted seeds have turned their later outcompeted by the slower, longer living
apex back towards their attachment point. trees.
involucre(Latin involucrum) A whorl of bracts lacerate Appearing as if irregularly torn or cleft.
subtending a flower cluster (as in the heads of laciniate Cut into narrow lobes or segments.
Compositae); in gymnosperms the whorl of lamina (Latin = blade, leaf) Here mostly used
scales (bud scales) subtending the strobilus in for relatively broad, flat leaves of conifers that
some species. would by most people not be described as nee-
isotype(Latin isotypus) A duplicate specimen of dles or scale leaves.
the holotype. laminar Of the leaf; also referring to organs
IUCN International Union for the Conservation homologous to leaves.
of Nature = The World Conservation Union. A laminated Divided into lobes (as of leaves).
non-governmental organisation (NGO) with lanateWoolly.
both government and non-government (e.g. lanceolate Of a shape reminiscent of a lance point,
academic) membership dedicated to the con- i.e. much longer than wide and with the great-
servation of natural resources, in particular bio- est width below the middle; commonly used to
logical species. Its Species Survival Commission describe the shape of a leaf.
(SSC) generates and updates the IUCN Red List laterals (lateral leaves) The paired, opposite scale
of Threatened Species; its categories and crite- leaves on a plagiotropic foliage branchlet (esp.
ria are used in this Handbook. in the family Cupressaceae) that are positioned
juvenile leaves Here the type of leaves that appears laterally, they flank and often partly enclose the
on seedlings, saplings, or immature trees, or differently shaped facials on either side.
laterite(Latin later = brick) A chemically weath- linear An elongated, narrow, flattened shape with
ered, earth-like type of rock rich in oxidized parallel or nearly parallel sides for all or most of
metals. its length.
lauraceous With alternate entire leaves character- lithosol An azonal shallow soil consisting of
istic of the family Lauraceae. imperfectly weathered rock-fragments.
layering The rooting of lower branches of a tree Ma Million years ago.
where they reach the soil; from these branches maquis A thick, scrubby (underbrush) vegetation
new plants may grow, which are for a time still of predominantly xerophytes (e.g. sclerophyl-
part of the old plant but eventually become lous shrubs) characteristic of Mediterranean
independent when the parent plant dies. shores and mountain slopes. [In New Caledonia,
1082 lectotype(Latin lectotypus) A specimen or illus- similar vegetation (but with very different plant
tration, designated from the original material species) growing on ultramafic soil rich in met-
as the nomenclatural type if no holotype was als is called maquis minier, as these substrates
indicated at the time of publication, or if it is are often mined to extract nickel.]
missing, or if it is found to belong to more than Malesia A floristic region of mainly islands in SE
one taxon. Asia, defined (mapped) in Flora Malesiana as
leg. ign.(Latin legitavit = collected by, ignotus = extending from the Isthmus of Kra on the Malay
unknown). The abbreviation is used here when Peninsula to the end of the main archipelago of
the collector of a herbarium specimen was not the Solomon Islands. The serial Flora Malesiana
stated. describes the plants of this region.
lenticels Usually more or less raised, often some- Massarts model A growth model in tree architec-
what corky spots on bark, of different shapes ture with a monopodial, erect stem with apical
but often horizontally elongate to linear, where dominance and branches placed in (pseudo)
gaseous exchange takes place. whorls at regular intervals; these branch again
lenticular Shaped like a lentil, i.e. with sides as in a and all or most branching is directed in a more
double-convex lens. or less horizontal plane (plagiotropic), by sup-
Life Zone A major altitudinal zone, character- pression of all but the laterally placed vegetative
ized by specific plant species and communities buds.
(and to a lesser degree by specific animal spe- median A position in the middle or central part
cies) in a mountain region; they are specifi- of an organ, e.g. a leaf.
cally recognized and named in Canada and the megagametophyteIn gymnosperms (conifers)
USA. the tissue inside the integument of an ovule or
lignified(Latin lignum = wood) Converted into seed coat of a seed, in which the fertilized gam-
wood; become woody. ete develops into an embryo.
lignite An organic deposit from plant remains in membranous Thin, rather soft and more or less
an intermediate phase of being transformed translucent.
from peat to bituminous black coal. meristem A formative plant tissue usually made
lignotuber An outgrowth or swelling on the basal up of small cells, capable of dividing continually
root system of a tree, forming excess wood and and giving rise to similar or differentiating cells.
capable of initiating new aerial stems with foli- mesa (Spanish = table) A flat topped mountain
age and reproductive organs. resulting from processes of erosion in ancient
ligulate Tongue-shaped; also (in description of sandstone or other sedimentary formations.
leaves) strap-shaped, resembling a ligulate mesic Characterized by or relating to a moderate
flower in Compositae. amount of moisture (see also mesophytic).
ligule(Latin ligula = small tongue) In the seed mesophyll Leaf tissue, consisting of spongy
cone scale of Araucaria the apical free part of parenchyma and palisade parenchyma (the
the seed scale, which is for the most part fused latter type not in all species) and filling the leaf
with the bract. between the hypodermis and the endodermis.
mesophytic Growing under moist conditions; also Groups that do not include all descendants are
of plants or plant communities growing under then described as paraphyletic.
medium conditions of moisture, without (pro- monopodial Having a single stem or trunk (bole),
longated) periods of draught (see also mesic). usually growing more or less straight up from
mesothermal Characterized by a moderate (cool) the ground.
temperature without pronounced daily or sea- monospecific Referring to a taxon of higher rank
sonal fluctuations. containing a single species (see also monotypic).
metamorphic Of a rock type that has been sub- monotypic Referring to a taxon containing only a
stantially altered due to high pressure and/or single taxon of lower rank, such as a genus with
heat generated by earth crust movements. only one species (see also monospecific).
micropyle The opening in an ovule through which montane Growing in, or being, the biogeographic 1083
pollen enters, in most conifers with the aid of a zone that is made up of relatively moist cool
pollination drop of fluid. upland slopes below timberline in mountainous
microsporangium(plural: microsporangia) A areas.
sac-like structure that contains the microspores, morphology The detailed study of form; in sys-
it is dehiscent so it can release the ripe spores. tematic research of taxa this involves investi-
In conifers this is usually termed pollen sac and gations into anatomy and development as well
it contains pollen. as comparison among individuals and taxa in
microsporophyll A (modified) leaf-like struc- order to understand characters and their states.
ture bearing the pollen sacs containing pollen mossy forest A high montane to subalpine forest
or male gametes (in gymnosperms); they are type in the tropics of SE Asia, characterized by a
borne on the male strobilus or pollen cone. low canopy of flat-topped crowns of trees, some
molecular analysis In the present context a phy- of them emergent, and draped with great quan-
logenetic analysis using DNA sequence data as tities of epiphytic mosses, lichens and ferns.
characters and cladistic methodology. mucronate Having a mucron, i.e. a narrow, abrupt
monoeciousIn gymnosperms this pertains to point which is usually only a continuation of the
female and male reproductive organs, com- midrib (or midvein).
monly cones, being present on the same plant. muskeg (word of Algonquin origin) A Sphagnum-
Plants with only one sex present are dioecious; built bog of northern latitudes in North America,
both conditions occur in conifers, with some often with scattered and stunted tree growth.
genera (or families) having one condition in all mutant An organism that obtained a genetic char-
species, but others more variable. acter change (mutation) not present in other
monograph In biology the comprehensive taxo- individuals of its species.
nomic description of a natural group of species, mutualism An ecological (or behavioral) relation-
usually a genus or a family, including all aspects ship between two species that is beneficial to
pertinent to the systematics and classification both.
of that group. mycelian Of the mycelium, the threadlike net-
monomorphic Of single form or shape (= 1char- work of hyphae of fungi.
acter state); all similar organs (e.g. leaves) mycelium A mass of filamentous hyphae, usually
have the same shape (compare dimorphic and underground or in wood, which forms the veg-
polymorphic). etative portion of a fungus.
monophyletic Adjective referring to a group of mycorrhiza The thread-like hyphae of certain
taxa; originally (Ernst Haeckel 1866) used in fungi that form intricate connections with the
the sense of sharing a unique common ancestor, roots of plants and live in symbiosis with these
but now almost universally used in the cladis- plants.
tic sense (Willi Hennig 1950, 1966), in which mycropyle The small opening at the apex of an
it refers to an ancestor and all its descendants. ovule, usually shaped somewhat like the neck of
[A more accurate term would be holophyletic.] a flask, through which pollen can enter.
natural group A group of taxa that is based on a numerical taxonomy(phenetics) Taxonomy
hypothesis of common descent from a nearest derived from the statistical analysis of variable
ancestor. data, i.e. the calculation of phenetic similarity
naviculate(Latin navicula = a boat) Boat shaped, between two taxa based on as many measurable
referring to the shape of open boats with nar- variables (character variations) as possible, usu-
rowing bow and stern and a more or less keeled ally by applying one of many available cluster-
bottom. ing algorithms and statistical probability tests.
neotype(Latin neotypus) A specimen chosen and Greater overall similarity is assumed to mean
designated as the type of a taxon in a situation closer relationship, thus directly informing the
where no original material is extant. classification.
1084 niveous white Snow-white, without any tinge of nutrient In the context of plants any of the basic
different (underlying) colours. elements such as nitrogen (N), potassium (K)
nom. cons.(Latin nomen conservandum = retained and phosphor (P) necessary for growth and
name) A botanical name invalid under the rules physiological processes.
of the Botanical Code (ICBN), but retained to n.v.(Latin non vidi) I have not seen it. Added to
avoid disadvantageous changes in the nomen- the citation of a (type) specimen one knows
clature by incorporation in the appended lists exists, but has not been able to see. Viewing a
of nomina conservanda (et rejicienda) published specimen can include seeing an image of it, e.g.
in the ICBN. on the Internet.
nom. illeg.(Latin nomen illegitimum = illegiti- obcordate Inversely heart-shaped.
mate name) A botanical name invalid under the obdeltoid Shaped like an inversed capital Greek
rules of the Botanical Code (ICBN), as specified letter delta (triangle).
under Art. 6.4 (Vienna Code, 2006). oblanceolate Considerably longer than broad,
nom. inval.(Latin nomen invalidum = invalid inversely lance-shaped (the widest part above
name) A botanical name that is not validly pub- the middle, compare lanceolate).
lished under the rules of the Botanical Code oblong Considerably longer than broad, with
(ICBN). Unlike an illegitimate name, it in fact nearly parallel sides.
does not exist for nomenclatural purposes unless obovate Inversely egg-shaped in outline
and until it has been validated under the rules. (2-dimensional).
nom. nud.(Latin nomen nudum) A botanical obovoidInversely egg-shaped (3-dimensional,
name without a formal description and there- compare pyriform).
fore invalidly published under the rules of the obtrullateInversely trullate, i.e. shaped like a
Botanical Code (ICBN). bricklayers trowel with the two short sides at
nom. ut. rej.(Latin nomen utique rejiciendum = the apex.
rejected name in any case) A botanical name obtuse Blunt, with a more or less rounded apex.
ruled as rejected under Art. 56 of the Botanical oligotraphent Adjective of a taxon, which predom-
Code (ICBN), listed in Appendix V and not to inantly grows in an oligotrophic environment.
be used in any case. oligotrophic Adjective of an environment (usu-
nothospecies (nothotaxon) A species (or taxon) ally a body of water), which is poor in dis-
which originated from the hybridization of two solved nutrients available to plants (compare
other species (or taxa) and is found to form eutrophic).
populations beyond the F1 generation with dis- ontogenetically (ontogeny) Referring to the pro-
tinct and more or less stable characters. cess of development or growth of an organism
nucleotide Any of the unit building blocks or or its parts from fertilized egg to adult.
bases of DNA and RNA, commonly repre- op. cit.(Latin opus citatus) Work cited. Referring
sented by the letters A (adenine), C (cytosine), here to the publication cited with a name of
G (guanine) and T (thymine); in RNA U (ura- a taxon under the species to which the text
cil) replaces T. In the DNA chain, A links with T applies; instead of to the list of references at the
and C with G, forming sequences of nucleotides. end of the Handbook.
orbicular Approximately circular in outline. pathogenic Referring to a pathogen, an agent or
orthotropic Growing of the stem (sometimes the organism that causes disease.
branches) in a vertical or near vertical direction, pectinate Arranged as the leaflets of a compound,
as opposed to the plagiotropic lateral branches pinnate leaf; with the needles (leaves) arranged
growing in a (near) horizontal direction. on both sides of the shoot, spreading sideways
ovate Egg-shaped in outline (mostly for plane and more or less in a plane (compare pinnate).
shapes). pedicellate With a small flower stalk (Latin pedi-
ovoidEgg-shaped. cellus); here used to describe the abruptly nar-
ovulate Referring to or having ovules. rowed base of seed scales.
ovule Seed bud, in gymnosperms (conifers) peduncle The stalk of a flower or inflorescence
including the embryonic stages of a seed up to (Latin pedunculus); usage of this term to desig- 1085
the event of fertilization, in which it is exposed nate the stalk of a conifer (female) cone is mor-
to air and develops specialized features to cap- phologically incorrect insofar as such a cone is
ture pollen. not an inflorescence, but a shoot with ovulifer-
ovuliferous Bearing the ovules (Latin ovulum); ous scales, but has been common practice and is
which after pollination and fertilization become maintained here for convenience (and likewise
the seeds. used for the stalk of male strobili).
palaeobotany A branch of botany (or geology) peltate Shaped more or less like a shield, with an
concerned with the study of fossil plants or their attachment more or less close to the centre.
remains. pendant Hanging down (nearly) vertically (see
palaeontology The science or study of fossil also pendulous).
organisms. pendulous Hanging down (nearly) vertically (see
palisade cells Perpendicular elongated parenchyma also pendant).
cells in the mesophyll near the surface of leaves. permafrost A permanently frozen layer of sedi-
palynology The science or study of pollen and ments or soil, of variable thickness, formed in
spores, both of recent and fossil plants. cold regions; of this layer the thin upper part on
paniculate Arranged in a panicle, i.e. a compound, the surface thaws in summer.
branching raceme. pentagonal Shaped like a pentagon, i.e. a polygon
papilliform Shaped like a nipple (Latin papilla). with five angles and five sides.
parallelism In an evolutionary context, the devel- perular scales The basal scales of buds which
opment of a similar character in two or more remain at the base of shoots or cones some time
lineages which do not share a nearest common after elongation or maturity.
ancestor; the character has therefore evolved petiolate Having a leaf-stalk (petiole); also: shaped
more than once and independently. like a petiole.
paraphyletic Adjective referring to a group of taxa phenotypic (phenotype) Relating to the anatomi-
that share a nearest common ancestor but, as cal and morphological appearance (characters)
a group, does not include all of that ancestors of an organism, in particular those determined
descendants (see also under monophyletic). by its genes (DNA, genotype) and a concomi-
paratype(Latin paratypus) A specimen cited in tant ontogeny.
the protologue that is neither the holotype photosynthesis Synthesis of chemical compounds
nor an isotype, nor one of the syntypes if two with the aid of radiant energy (light), resulting
or more specimens were simultaneously desig- in the formation of carbohydrates (assimilates)
nated as types. from carbon dioxide and water in chlorophyll-
parenchyma A tissue of higher plants composed containing plant leaves (see also under chlorophyll).
of thin-walled cells; especially such tissue as the phyletic Referring to both common ancestry rela-
pith and mesophyll. tionships (phylogeny) and ancestor/descendant
passim(Latin passus = scattered) After a page relationships (diversification of characters /
number, indicating that the name (subject) sub- character states through a hypothetical evo-
sequently appeared on several other pages. lutionary lineage resolved in the phylogeny)
(compare phylogenetic).
phylloclade (phyllode) (Greek phyllos = leaf, cla- multi-layered wall and is dispersed to meet an
dos = branch) A leaf-like branch; in the genus ovule for fertilization.
Phyllocladus true leaves are rudimentary and pollen sac (microsporangium) A variously shaped
foliage branchlets are strongly flattened, func- sac-like vessel containing pollen and which is
tioning as leaves. borne on the microsporophyll of a pollen cone
phyllotaxis The arrangement of leaves on a shoot in conifers.
or stem and their position in relation to each pollination The transfer of pollen from a stamen
other. (in angiosperms) to an ovule; in gymnosperms
phylogeneticReferring to common ancestry the pollen descends directly on the nucellus and
relationships between extant species or taxa germinates; the pollen tube begins to grow and
1086 of higher ranks (see phylogeny and compare penetrates the nucellus.
phyletic). polymorphic Of various forms or shapes; similar
phylogeny A hypothesis of common ancestry rela- organs on the same plant appear in > 2char-
tionships of taxa, usually depicted as a branch- acter states, e.g. variable leaf forms (compare
ing diagram or tree (Tree of Life) showing dimorphic and monomorphic).
branching relationships from a hypothetical polyphyletic Referring to a group of taxa that do
common ancestor to multiple descendants. not share a nearest common ancestor.
Cladograms, although really only showing rela- polytomy In a cladogram referring to lineages
tionships of characters and character states, are (clades) with unresolved relationships; such lin-
usually interpreted as phylogenies by inserting eages all arise from a single node.
taxa on the (terminal) branches, inferring their population The sum total of the individuals of a
phylogenetic relationships from the branching species (or lower taxon) which inhabit a certain
pattern (topology) of the cladogram. area and are expected to interbreed (exchange
phytogeography The study of natural plant (usu- genes).
ally species) distribution and its causes. p.p.(Latin pro parte = in part) adjective to a name
pilose Having straight, soft and spreading hairs. indicating that only a certain part (including
pinetum (plural: pineta) A park or garden dedi- or excluding the type) of the original material
cated to a living collection of planted conifers. mentioned in the protologue is (to be) con-
pinnate Having the arrangement of a feather, nected with that name.
with a single rachis from which leaflets arise on primary branches The branches that appear more
either side (compare pectinate). or less rhythmically during the growth of a tree,
plagiotropic Growing of (lateral) branches in a from first to highest order. Secondary branches
horizontal or near horizontal direction, away are the result of reiteration. The distinction
from the (near) vertical (orthotropic) stem between the two categories becomes increas-
or first order branches. ingly blurred and often difficult to determine in
planation The transformation through evolution old trees, in which much of the main branches
of a more or less terete shape of something in may represent reiteration.
an ancestor into one that becomes flattened and primary leaves The first leaves of a young plant,
wider in its descendants. other than the cotyledons. In some genera (or
plate tectonics The formation and dynamics of the species) these leaves differ markedly from later,
plates into which the crust of the Earth is divided. adult types of leaves, e.g. in Pinus (cataphylls),
podzolic (podzolized, Russian podzol = under in others the differences are gradual, e.g. in
ashes) Referring to an infertile, sandy and Podocarpus (compare secondary leaves).
acidic soil with minerals leached from its sur- primary (bud) scales The outer scales, which are
face layers and deposited in a lower stratum. often much longer than the inner (second-
The leached part of the profile is ash-grey. ary) scales in foliage branches of the genus
pollen (singular and plural) The (near) micro- Podocarpus.
scopic unit or grain in seed plants that contains prismatic Shaped like a prism, i.e. with polygonal
the male gamete enclosed by a hard and usually faces lying in parallel planes.
proliferation (in conifers) The development of selves (becoming orthotropic) cause repetition
a foliage shoot from a normally determinate of the first order branching in (pseudo-)whorls.
organ, e.g. a seed cone. receptacle A fleshy or succulent structure subtend-
protologue(Greek protos =first; logos = discourse) ing a free standing seed in Podocarpus and some
Everything associated with a name at its first other genera of the Podocarpaceae; it is formed
valid publication, i.e. diagnosis, description, from all remnants of the seed cone after fertili-
illustrations, references, synonymy, geographi- sation of usually a single egg (ovule) developing
cal data, citation of specimens, discussion, and into the seed.
comments. recurved Curved backward (compare decurved).
provenance Information about the source of a refugium (plural: refugia) A geographical area
plant or its seed used in forestry or horticulture. into which one or more species have retreated 1087
proximal Occurring nearest the base or axis (com- (or where they remained) from a much wider
pare distal). distribution in the past.
pruinose Covered with a coarse, whitish, waxy reiteration(Latin reiterare = to say or do repeat-
bloom, more prominent than if glaucous. edly) The secondary initiation of branching
pseudo-whorl An arrangement that is seemingly from a primary branching system (or from the
forming a whorl, but is in fact helical on a very trunk) of a tree, activating dormant buds, usu-
short axis. ally as a response to damage.
puberulentMinutely pubescent, i.e. covered with rendzina A dark greyish brown intrazonal soil
very small, soft hairs. developed in usually grassy regions of high to
pubescent Covered with soft, short hairs. moderate humidity from soft calcareous marl
pulverulent Being or looking dusty or powdery or chalk.
(from almost invisible short hairs). reniform Kidney-shaped; much wider than long.
pulvinus (plural: pulvini) Small, peg-like projec- repand With an undulating or wavy margin.
tion on the shoot supporting the leaf (needle) resin duct (resin canal) A tubular intercellular
in several genera especially of Pinaceae. space, especially in gymnosperms, that is lined
pungent Ending in a rigid, sharp point or prickle. with cells which secrete resin.
pustulate Having pustules, i.e. low projections like resin vesicle A small cavity or bladder filled with
a blister or pimple. resin.
pyriform Shaped like a pear, i.e. with the broadest reticulate Forming a network of connections or
part near the apex (compare obovoid). relationships between distinct entities.
quadrate In fours, here pertaining to the phyllotaxis, retuse Notched shallowly at a rounded apex.
with four leaves originating at exactly the same revolute Rolled downward or backward.
distance on the shoot. If one of the opposite pairs rheophyte A plant that completes its life cycle in
is slightly higer, the arrangement is decussate. streams, but is not aquatic; a rheophyte germi-
quadrate-rhombic Shaped between quadrangular nates out of water when the stream falls tem-
and rhombic (compare rhombic). porarily dry or recedes from normally higher
raceme An inflorescence with the oldest flowers levels.
(pollen cones in conifers) the most proximal rhombic Having the shape of a rhombus, i.e.
and a potentially continuously growing apex. an equilateral parallelogram usually having
racemose In the form of a raceme. oblique angles (compare rhomboid).
rachis The axis (homologous to a shoot) of the rhombic-orbicular Shaped between rhombic and
cone of a conifer, from which bracts, microspo- orbicular.
rophylls or seed scales arise. rhomboid shaped like a parallelogram in which
Rauhs model A growth model in tree architec- the angles are oblique and adjacent sides are
ture with a monopodial, erect stem with apical unequal (compare rhombic).
dominance and branches placed in (pseudo-) riparian Growing on the banks of rivers, subject to
whorls at regular intervals; these and secondary flooding, but not permanently in water.
branches are assurgent and by erecting them-
RNA Ribonucleic acid. In its various forms, RNA septal(Latin septum = enclosure) Here a position
acts as the intermediary by which the heredi- of the resin canals in the leaves wedged between
tary code of DNA is converted into proteins the endodermis (enclosing the inner vascular
(compare DNA). cylinder) and the hypodermis.
rostrate Shaped like a beak (Latin rostrum). sequencing(of DNA) A technique using bio-
ruderal Referring to plants or vegetation growing chemical methods to determine the sequence
in and adapted to continuously or repeatedly of the four nucleotides (the genetic code) in
disturbed sites (dynamic habitats). a particular section of the DNA strand of an
saccate With bladders (Latin sacci); some types of organism. These sequences are copied through
pollen in conifers have 2 (rarely more) hollow generations, but small changes (mutations)
1088 bladders, presumably to aid buoyancy. occur from time to time, which are again copied
sapwood The outer section of the wood of a tree, and so serve as markers of heredity. They can be
with living wood cells through which water is identified and scored as derived characters for a
transported. cladistic analysis.
scarious Thin, dry and membranous, not green sere A successional sequence of plant associations
(herbaceous). or communities that replace each other over
scion A vegetative shoot cut from a plant and time at a given site, usually going from pio-
caused to produce roots or grafted onto a differ- neer stages to later stages in which an ecologi-
ent rootstock. cal equilibrium is reached. The stages are called
sclereids(Greek skleros = hard) Inclusions in the seral to emphasize their transitory nature.
mesophyll or below the surface of leaves con- serotinous(Latin sero = late) Late to appear or
sisting of stone cells, i.e. dense and hard bodies flower; in conifer seed cones pertaining to a
often with a random position and distribution. delayed opening, usually associated with persis-
sclerified Hardened by transformation or addition tence on branches; both are adaptations to fire.
of cells with lignified walls. serrate Saw-toothed, the sharp teeth pointing
sclerophyllous With hard leaves; leaves with a forward.
tough texture, mainly from dermal cells with serrulateFinely serrate.
lignified walls. sessile(Latin sessilis = sitting, attached) Without
secondary leaves Leaves of the adult type; in coni- a peduncle or stalk, or in the case of conifer
fers these may appear soon on young plants, or cones, with a very short one which is invisible
they may be delayed for much longer and they under the basal scales so that the cone appears
are the forms that are usually associated with to be stalkless.sister groupA clade in a clado-
fertile branches and/or appear in the canopy gram that is nearest in relationship to another
of mature trees or shrubs (compare primary clade, or a taxon represented by such a clade.
leaves). The term usually (but not necessarily) refers to
seed scale Appendage in a conifer seed cone (situ- the clade below the next one in a cladogram.
ated in the axil of a bract) bearing one or more sister (group or taxon). In cladistic terminology a
seeds. group or a taxon that is a clade next to another
sensu lato (Latin = in a broad sense) In taxon- clade, of which it is the sister clade; within a par-
omy it follows a taxon name to indicate that it ticular cladogram these two are more closely
includes another taxon at the same rank, which related to each other phylogenetically than they
some may recognize as distinct (abbreviated are to any other clades (groups) represented.
ass.l.). The opposite is sensu stricto. s. l.(Latin sensu lato) In a broad sense; used in
sensu stricto (Latin in a strict or narrow sense). addition to a taxonomic name to differentiate it
In taxonomy it follows a taxon name to indi- from the use of that name in a strict sense (see
cate that it excludes another taxon at the same sensu lato).
rank, which some may consider as synonymous s.n.(Latin sine numero) Without a number. If col-
(abbreviated as s. str.). The opposite is sensu lectors of botanical specimens did not give a
lato. number to them, such collections are cited with
the name of the collector and s.n. immediately subalpine Referring to, or growing in the biogeo-
following. graphic zone between the montane zone and
s. str.(Latin sensu stricto) In a strict or narrow the tree limit in high mountains; it is the alti-
sense (see sensu stricto). tude of the highest growing tree species in a
spathulateSpoon-shaped. given area.
speciation To differentiate into new biological substrate The base on which an organism lives; for
species through evolution, involving genetic most vascular plants this is the soil.
separation of populations and divergence of subtending Occupying an adjacent and usually
characters (character states) through time. lower position to and often (partly) enclosing
species complex Term used to describe a group of an organ.
closely related species, within which group spe- subtereteAlmost terete, the shoot may be very 1089
ciation takes place (variation leading to genetic slightly angular or ridged.
separation) and/or which is often character- subulate Awl-shaped; linear and tapering to a fine
ized by hybrids (occurrence of hybrid-swarms, point.
reticulate relationships between taxa). succession In ecology, the gradual and successive
speciose Counting or having many species. replacement of plant species by others in one
spicate As in a spike, spike-like. locality due to development of the vegetation
spike An inflorescence consisting of an axis and from a pioneer phase to a climax phase.
numerous flowers (pollen cones in conifers) succulent Of a plant: having fleshy tissues adapted
arranged on it (see also raceme). to conserve and store moisture (like the cac-
spinescent Shaped like a spine or thorn (compare tuses or cacti).
pungent). symbiosis A mutually beneficial physiological
spiral Arranged or shaped in an outward going, relationship between two or more different spe-
circular fashion. On shoots, this will be an cies, with give and take in equilibrium in the
upward spiral and is more accurately termed sense of an evolutionary stable strategy (ESS).
helical, but spirally arranged leaves is often sympatric Occurring in (generally) the same area
used, including in this book. or with partly overlapping areas, but without
sporophyll Literally a leaf bearing spores (as in loss of genetic identity by interbreeding.
ferns); in gymnosperms it refers to leaf-like syntype(Latin syntypus) Any specimen cited in
appendages bearing male (= microsporophyll) the protologue when no holotype was desig-
or female (= macrosporophyll) reproductive nated, or any of two or more specimens simul-
organs: pollen (in pollen sacs) or ovules. taneously designated as types.
squarroseWith stiff or rigid branches or systematics The science of the diversity of and the
protrusions. relationships between taxa based on evolution-
stele An axial cylinder of tissue in which the vas- ary principles, including (or sometimes seen as
cular tissue is developed; its outer layer of cells synonymous with) taxonomy.
forms the endodermis. synapomorphy (plural: synapomorphies) Shared
stomata (sing. stoma) Breathing pores or apertures derived character(s); i.e. a character originated
in the epidermis, surrounded by two guard cells in a common ancestor and inherited unal-
(and subsidiary cells), leading into an intercellu- tered by its descendants; it could be lost subse-
lar space communicating with the internal tissue. quently in some descendants and transformed
striated Marked with (longitudinal) lines or into a different character in others (see also
streaks. homologous).
strobilus(Greek strobilos = pine cone; plural: taiga (Russian = forest) Northern coniferous for-
strobili) The technical term for the reproduc- est between tundra in the north and steppe in
tive unit of a gymnosperm that bears male or the south, particularly in Asia.
female organs, regardless of whether it forms a talus A slope formed especially by accumulation
distinct cone or not. of rock debris from rock formations above it.
taxon (plural: taxa) Any group of organisms that transverse Made at right angles to the anterior-
is recognized at any of the ranks of classifica- posterior axis of a body or structure; also set
tion used by taxonomists, such as family, genus, crosswise.
species. transversely rhombic Having the shape of a
taxonomy The science of classification of organ- rhombus (see rhombic), which is transversely
isms and the identification and naming of taxa. widened (see transverse).
TDWGInternational Working Group on triangulate Appearing as a triangle but not really
Taxonomic Databases for Plant Sciences shaped so.
(TDWG). The codes used in this Handbook are tridentate Three-toothed; in the bract scales of
for politically defined geographical areas (coun- cones of Pinaceae with a short, pointed central
1090 tries, provinces etc.) according to the World lobe flanked by two short, pointed lateral lobes
Geographical Scheme for Recording Plant (see also trilobate).
Distributions, compiled by R. K. Brummitt trigonal Having three sides or faces derived from
(2001). See with this reference for access. a triangular base.
terete Approximately cylindrical. trilobate Three-lobed; in the bract scales of cones
terminal Referring to the position of an organ or of Pinaceae with a long or rounded central lobe
structure at the tip of a shoot. flanked by two long or rounded lateral lobes
ternate Occurring in threes; leaves are ternate (see also tridentate).
only if all 3 leaves originate at exactly the same trimerous In threes; i.e. made up of three separate,
distance on the shoot, otherwise they are more free parts (compare tripartite).
likely to be alternate. tripartite Made up of three separate but connected
terpenoid (German: Turpentine) Referring to parts (compare trimerous).
chemical compounds based on a five-carbon trisaccate(in pollen) Having three sacci or air
atom structure and present in conifer resins, bladders (compare bisaccate).
also known as terpenes. trullate Shaped like a bricklayers trowel.
tesselate elements or pieces laid closely together as truncate Ending abruptly, as if cut off transversely
in a mosaic or like paving stones, joined only at (but usually with rounded corners).
the margins (compare imbricate). tussock In grassland, clumps formed by the con-
tetragonal Having four sides or faces derived from stituent grasses that are raised above the gen-
a quadrangular base. eral surface, such grassland is very uneven as a
tetrastichous Arranged to forming 4 apparent result.
rows from a helical leaf attachment on the type (type specimen) A specimen designated as
shoot. the type of the name of a species or taxon with
tomentose Densely covered with short, woolly lower rank, fixing the application of that name
hairs. by including its characters. The circumscrip-
tracheid A dead cell in the wood of gymnosperms tion of a taxon can be widened or narrowed,
characterized by lateral pits in the cell walls con- but always must include that of the type speci-
necting the lumen of one tracheid with that of men if it is to retain its name. Similar principles
another, allowing fluid transport; tracheids are apply at higher ranks, where e.g. a species name
structurally distinct from the equivalent cells in becomes the type of a genus, but in fact refers
angiosperms (vessels). to the type specimen of that species name.
translucent Of a substance or thing with a colour, ultrabasic Referring to rock or soil with a high pH
density, or thickness that allows some light to rich in metallic minerals (also ultramafic).
penetrate, but insufficient to be seen through ultramafic Referring to rock or soil poor in silica,
(compare transparent). but extremely rich in iron and magnesium min-
transparent Of a substance or thing with a colour, erals and with a high pH value (also ultrabasic).
density, or thickness that allows it to be seen umbellate Borne in umbels, i.e. a (flower) cluster
through (compare translucent). in which the pedicels or stalks (branches) arise
from a common point.
umbilicate Depressed like a navel. originally referred to a method of analysis in
umbo (orig. Latin = the boss of a shield) In pine which area cladograms were substituted for
cones it is a prominence on the apophysis of the taxon cladograms. This concept is intimately
scales, often armed with a prickle or spine. connected with cladistic methodology, under
undulate Having a wavy surface, edge, or mark- the assumption that distribution patterns and
ings (see also repand). speciation are caused by the same historic
utriculate Inflated, bladder-like. events. It ignores (later) distribution of taxa
valvate Parts connecting or touching with their caused by dispersal of organisms.
edges (as in the two shells of an oyster), not vicariant(Latin vicarius = substitute) A taxon
overlapping (compare imbricate). occupying a similar niche as a different, but
variegated Variably coloured in foliage or leaves, related taxon occurring in a separate area (vicar- 1091
usually occurring in cultivars from a partial iants are allopatric). The two taxa are inferred to
deficiency in the amount of chlorophyll in the have derived from an (extinct) common ances-
leaves, causing a yellowish hue contrasting with tor and their separation the result of the emer-
normal green. gence of an ecological barrier between them.
vascular bundle A unit of the vascular system of whip shoot A relatively fast growing, leading foli-
a vascular plant, consisting of vessels and sieve age branchlet; due to its faster growth scale
tubes, together with parenchyma cells and leaves, which grow with it, are longer and often
fibres; it has a function in transportation of also wider than those on short, lateral branchlets.
water, nutrients and assimilates. winter bud A terminal bud, which in conifers is
vegetative bud (vegetative apex) A bud which covered with scales or cataphylls which can be
gives rise to the vegetative organs, i.e. the shoot, resinous or non-resinous, and which contains
dwarf shoots and needles or leaves. In the the young shoot of which the internodes are not
descriptions of this Handbook usually the ter- yet elongated; it will start growing at the begin-
minal buds at the end of branches are the only ning of the spring season (see also vegetative
ones described (see also winter bud). bud).
venation The number and distribution (pattern) xeromorphic Structurally adapted for life and
of veins in a leaf. growth with a limited water supply, especially a
verrucose Covered in numerous small, wart-like morphology limiting transpiration (in gymno-
elevations (verrucae). sperms) or providing for the storage of water;
vessel (in wood) A cell in the wood of angio- such plants are xerophytes.
sperms (and some gymnosperms) with closed xerophyte A plant structurally adapted for life
lateral walls and distal openings allowing fluid and growth with a limited water supply; usu-
transport (compare tracheid). ally by means of xeromorphic adaptations that
vicariance(Latin vicarius = substitute) In bioge- allow water storage and ensure limitation of
ography referring to the distribution of taxa transpiration.
explained by the history of the separation of zoochory The dispersal of plant propagules (e.g.
the areas in which these taxa occur. The term seeds) by animals.
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L I S T S O F I L LU S T R AT IO N S
121. Falcatifolium falciforme trees in the Crocker 125. Fitzroya cupressoides in the N. P. Alerce Andino,
Range, Borneo Chile (photo P. Woltz)
122. Falcatifolium falciforme flushing leaves 126. Fitzroya cupressoides foliage and seed cones
123. Falcatifolium falciforme seedling at Frasers Hill, 127. Fokienia hodginsii foliage and cones
Malaysia 128. Glyptostrobus pensilis pollen cones and seed
124. Falcatifolium taxoides on Mt. Pani, New cones (photo D. White)
Caledonia 129. Halocarpus bidwillii in North Island, New
Zealand
Halocarpus bidwillii foliage
130. 152. Keteleeria davidiana var. davidiana seed cone
Halocarpus biformis foliage
131. 153. Lagarostrobos franklinii tree at Riveaux Creek,
Juniperus californica in Anza Borrego Desert
132. Tasmania, Australia
State Park, California, USA 154. Lagarostrobos franklinii foliage and pollen
Juniperus californica seed cones
133. cones
Juniperus chinensis var. sargentii foliage and
134. 155. Larix decidua var. decidua in the Alps,
seed cones Switzerland
Juniperus communis var. communis foliage and
135. 156. Larix decidua var. decidua bark
seed cones 157. Larix decidua var. decidua seed cones
Juniperus communis var. saxatilis in Mt. Rainier
136. 158. Larix gmelinii var. principis-rupprechtii seed
1108 N. P., Washington, USA cones
137. Juniperus deppeana var. deppeana in Puebla, 159. Larix griffithii var. griffithii seed cone
Mexico 160. Larix kaempferi seed cones
138. Juniperus deppeana var. deppeana trunk with 161. Larix lyallii in the Wenatchee Mts., Washington,
bark USA
139. Juniperus flaccida var. flaccida tree in Oaxaca, 162. Lepidothamnus fonkii in Chile (photo
Mexico M. Gardner)
140. Juniperus occidentalis var. australis tree in the 163. Libocedrus bidwillii in North Island, New
Sierra Nevada, California, USA Zealand
141. Juniperus oxycedrus subsp. macrocarpa foliage 164. Libocedrus bidwillii foliage and seed cones
and seed cones 165. Libocedrus plumosa foliage
142. Juniperus phoenicea subsp. phoenicea at Cape 166. Manoao colensoi seed cones (photo
St. Vincent, Portugal B. P. J. Molloy)
143. Juniperus phoenicea subsp. phoenicea foliage 167. Metasequoia glyptostroboides bark
and cones 168. Metasequoia glyptostroboides seed cones
144. Juniperus pseudosabina in the Alaj Mountains, 169. Microbiota decussata foliage
Kirgyzstan 170. Microcachrys tetragona foliage and seed
145. Juniperus pseudosabina seed cones in cones
Kirgyzstan 171. Nageia fleuryi leaves and seed cones (photo
146. Juniperus sabina var. sabina in Kirgyzstan L. Averyanov)
147. Juniperus semiglobosa in Kirgyzstan 172. Nageia nagi flushing leaves
148. Juniperus semiglobosa foliage and seed cones 173. Nageia wallichiana tree in Viet Nam (photo
149. Juniperus semiglobosa pollen cones L. Averyanov)
150. Juniperus virginiana var. virginiana tree in 174. Nageia wallichiana leaves and seed cones
North Carolina, USA (photo L. Averyanov)
151. Keteleeria davidiana var. davidiana foliage and 175. Neocallitropsis pancheri in New Caledonia
seed cones 176. Neocallitropsis pancheri foliage
240. Pinus pinaster subsp. pinaster pollen cones 250. Pinus taiwanensis var. taiwanensis in Hehuan
241. Pinus pinaster subsp. pinaster seed cones of two Shan, Taiwan
ages 251. Pinus virginiana seed cones
242. Pinus pinea in Algarve, Portugal 252. Pinus wallichiana var. wallichiana seed cones
243. Pinus pinea seed cone 253. Platycladus orientalis foliage and seed cones
244. Pinus ponderosa var. ponderosa pollen cones (photo D. Mabberley)
245. Pinus pungens seed cones 254. Platycladus orientalis seed cones
246. Pinus quadrifolia in California, USA 255. Podocarpus acutifolius foliage and pollen cones
247. Pinus rzedowskii tree in Michoacn, Mexico 256. Podocarpus brassii var. brassii seed cone (photo
248. Pinus rzedowskii seed cones T. Waters)
249. Pinus sylvestris var. sylvestris trees 257. Podocarpus brevifolius on the Mesilau River,
Mt. Kinabalu, Borneo
258. Podocarpus brevifolius leaves 280. Podocarpus nivalis in North Island, New
259. Podocarpus chingianus foliage and seed cone Zealand
260. Podocarpus costalis foliage 281. Podocarpus nivalis foliage and seed cones
261. Podocarpus cunninghamiii foliage and seed 282. Podocarpus novae-caledoniae in New Caledonia
cones 283. Podocarpus novae-caledoniae foliage and young
262. Podocarpus dispermus leaves seed cones
263. Podocarpus elatus foliage 284. Podocarpus nubigenus leaves (upperside)
264. Podocarpus elatus seed cone 285. Podocarpus nubigenus leaves (underside)
265. Podocarpus grayae small sapling at Cape 286. Podocarpus polystachyus pollen cones
Tribulation, Queensland, Australia 287. Podocarpus rumphii seed cones
1110 266. Podocarpus grayae tree at Lake Eacham, 288. Podocarpus salignus foliage and young seed
Queensland, Australia cones
267. Podocarpus grayae trunk, Herberton Range, 289. Podocarpus spinulosus on North Stradbroke
Queensland, Australia Island, Queensland, Australia
268. Podocarpus henkelii leaves 290. Podocarpus spinulosus unripe seed cone (photo
269. Podocarpus laubenfelsii seedling on Mt. G. Garruthers
Kinabalu, Borneo 291. Podocarpus spinulosus ripe seed cones (photo
270. Podocarpus latifolius in the Drakensberg, South G. Garruthers)
Africa (photo J. Grimshaw) 292. Podocarpus sprucei foliage buds (photo P. Cribb
271. Podocarpus lawrencei seed cones 1425)
272. Podocarpus macrophyllus var. macrophyllus 293. Podocarpus totara tree in North Island, New
foliage Zealand
273. Podocarpus macrophyllus var. macrophyllus 294. Podocarpus totara bark
foliage and pollen cones 295. Podocarpus totara foliage and pollen cones
274. Podocarpus matudae leaves and young seed 296. Prumnopitys andina foliage and pollen cones
cone 297. Prumnopitys andina seed cones
275. Podocarpus milanjianus on Mt. Elgon, Uganda 298. Prumnopitys ferruginoides foliage on Mt. Mou,
(photo D. L. Roberts) New Caledonia
276. Podocarpus nakaii flushing leaves 299. Prumnopitys ladei trunk on Mt. Lewis,
277. Podocarpus nakaii foliage and seed cones Queensland, Australia
278. Podocarpus neriifolius var. neriifolius in Papua 300. Prumnopitys ladei foliage
New Guinea (photo T. Utteridge) 301. Pseudolarix amabilis pollen cones
279. Podocarpus neriifolius var. neriifolius seed cones 302. Pseudolarix amabilis seed cones
(photo T. Utteridge) 303. Pseudotaxus chienii foliage and buds of seed
cones
304. Pseudotsuga japonica foliage and young seed 310. Pseudotsuga sinensis var. sinensis fallen seed
cone cones
305. Pseudotsuga japonica seed cones 311. Retrophyllum comptonii young tree on
306. Pseudotsuga menziesii tree in Mt. Rainier N. P., Mt. Pani, New Caledonia
Washington, USA Retrophyllum comptonii foliage and seed (photo
312.
307. Pseudotsuga menziesii giant trunk in Olympic M. Gardner)
N. P., Washington, USA Retrophyllum minus in Rivire des Lacs, New
313.
308. Pseudotsuga menziesii var. menziesii seed Caledonia
cones Retrophyllum minus foliage and seed cones
314.
309. Pseudotsuga sinensis var. sinensis trees in Taroko Saxegothaea conspicua tree (photo C. N. Page)
315.
N. P., Taiwan
316. Saxegothaea conspicua foliage with pollen 339. Tetraclinis articulata seed cones (photo
cones and seed cones M. Gardner)
317. Sciadopitys verticillata foliage and pollen cone 340. Thuja koraiensis in Korea (photo Y. S. Kim)
buds 341. Thuja koraiensis foliage branch (upperside)
318. Sciadopitys verticillata seed cone (photo 342. Thuja koraiensis foliage branch (underside)
C. N. Page) 343. Thuja plicata foliage and seed cones
319. Sequoia sempervirens at Bull Creek, California, 344. Thuja sutchuenensis in Daba Shan, China
USA (photo R. Van Pelt) ( Bedgebury Pinetum)
320. Sequoia sempervirens seed cones 345. Thujopsis dolabrata var. dolabrata foliage
321. Sequoiadendron giganteum in California, USA (upperside)
(photo E. Parker) 346. Thujopsis dolabrata var. dolabrata foliage 1111
322. Sequoiadendron giganteum pollen cones and (underside)
seed cones 347. Torreya californica foliage with pollen cones
323. Sundacarpus amarus trunk and bark, Lake 348. Torreya californica foliage with seeds
Barrine, Queensland, Australia 349. Torreya nucifera foliage with seeds
324. Sundacarpus amarus foliage 350. Tsuga chinensis var. chinensis in the mountains
325. Sundacarpus amarus pollen cones of Taiwan
326. Taiwania cryptomerioides in Yunnan, China 351. Tsuga chinensis var. chinensis foliage and seed
(photo D. Long) cones
327. Taiwania cryptomerioides tree (photo P. Thomas) 352. Tsuga forrestii foliage and seed cones
328. Taiwania cryptomerioides trunk in Taiwan 353. Tsuga heterophylla on a nurse log in Olympic
329. Taxodium distichum swamp forest in North N. P., Washington, USA
Carolina, USA 354. Tsuga mertensiana var. mertensiana foliage and
330. Taxodium distichum var. distichum foliage and seed cones
seed cones 355. Widdringtonia cedarbergensis seed cones
331. Taxodium mucronatum in Oaxaca, Mexico 356. Widdringtonia nodiflora seed cones
332. Taxus baccata ancient tree on ruined wall of 357. Widdringtonia whytei on Mt. Mulanje, Malawi
Waverley Abbey, England (photo P. Cribb)
333. Taxus baccata foliage and pollen cones 358. Wollemia nobilis trees in Wollemi N. P., New
334. Taxus baccata foliage and seeds South Wales, Australia (photo J. Plaza)
335. Taxus brevifolia in the Wenatchee Mts., 359. Wollemia nobilis bud and foliage
Washington, USA 360. Wollemia nobilis foliage and pollen cones
336. Taxus brevifolia foliage and seeds 361. Xanthocyparis nootkatensis trunk in Olympic
337. Taxus cuspidata var. cuspidata in Korea (photo N. P., Washington, USA
Y. S. Kim) 362. Xanthocyparis vietnamensis foliage
338. Taxus cuspidata var. cuspidata foliage and seeds
I N D E X T O B O TA N IC A L NA M E S O F C O N I F E R S
Nageia Gaertn. 20, 28, 42, 53, 175, 334, 378, 536, P
539, 870
Nageia amara (Blume) F. Muell. 983 Pachylepis Brongn.1056
Nageia argotaenia (Hance) Kuntze 175 Pachylepis cupressoides (L.) Brongn. 1056
Nageia comptonii (J. T. Buchholz) de Laub. 967 Papuacedrus H. L. Li 38, 48, 50, 330, 552, 553
Nageia falcata (Thunb.) Kuntze 142 Papuacedrus arfakensis (Gibbs) H. L. Li 554
Nageia fleuryi (Hickel) de Laub. 180, 258, 277, Papuacedrus papuana (F. Muell.) H. L. Li 21, 329,
472, 537, 1008 330, 333, 337, 552, 643, 835, 852
Nageia formosensis (Dummer) C. N. Page 540 Papuacedrus papuana (F. Muell.) H. L. Li var.
Nageia latifolia (Wall.) Gordon 541 arfakensis (Gibbs) R. J. Johns 552, 554
Nageia mannii (Hook. f.) Kuntze 145 Papuacedrus papuana (F. Muell.) H. L. Li var.
Nageia maxima (de Laub.) de Laub. 538, 890 papuana 552, 553, 643
Nageia minor Carrire969 Parasitaxus de Laub. 43, 53, 54, 347, 555
Nageia motleyi (Parl.) de Laub. 539 Parasitaxus usta (Vieill.) de Laub. 20, 381, 555,
Nageia nagi (Thunb.) Kuntze 269, 472, 536, 540 556, 643
Nageia nagi (Thunb.) Kuntze var. formosensis Parolinia Endl.1056
(Dummer) Silba 540 Pherosphaera W. Archer 43, 53, 54, 557
Nageia nagi (Thunb.) Kuntze var. koshuensis Pherosphaera fitzgeraldii (F. Muell.) Hook. f.
(Kaneh.) D. Z. Fu 540 557, 558
Nageia nankoensis (Hayata) R. R. Mill 540 Pherosphaera hookeriana Hook. f. 534
Nageia pancheri (Brongn. & Gris) Kuntze 133 Pherosphaera hookeriana W. Archer 223, 225,
Nageia piresii (Silba) de Laub. 970 374, 534, 557, 558, 559, 561, 643, 879
Nageia rospigliosii (Pilg.) de Laub. 971 Pherosphaera niphophila (J. Garden & L. A. S.
Nageia usta (Vieill.) Kuntze 555 Johnson) Florin 558
Nageia vitiensis (Seem.) Kuntze 972 Phyllocladaceae Bessey 18, 21, 24, 26, 35, 39, 45,
Nageia wallichiana (Presl) Kuntze 17, 24, 155, 181, 50, 51, 560
182, 277, 352, 377, 378, 472, 541, 542, 1017, 1068 Phyllocladus Rich. ex Mirb. 39, 50, 51, 358, 379,
Neocallitropsis Florin 39, 48, 50, 543 560, 563, 564, 565, 837
Neocallitropsis araucarioides (R. H. Compton) Phyllocladus alpinus Hook. f. 565, 566
Florin543 Phyllocladus aspleniifolius (Labill.) Hook. f. 223,
Neocallitropsis pancheri (Carrire) de Laub. 249, 225, 374, 560, 561, 643
350, 472, 522, 527, 543 Phyllocladus aspleniifolius (Labill.) Hook. f. var.
Nothocallitris A. V. Bobrov & Melikyan 249 alpinus (Hook. f.) H. Keng 566
Phyllocladus billardieri Mirb.560 Picea asperata Mast. 84, 118, 126, 439, 567, 574,
Phyllocladus glaucus Kirk 563, 564 576, 578, 583, 605, 612, 614, 623
Phyllocladus hypophyllus Hook. f. 334, 336, 359, Picea asperata Mast. var. asperata 569, 576, 613
360, 553, 562, 563, 644, 843, 852, 863, 909, 984 Picea asperata Mast. var. aurantiaca (Mast.) Boom
Phyllocladus toatoa Molloy 563, 564, 566 577
Phyllocladus trichomanoides D. Don 152, 332, Picea asperata Mast. var. notabilis Rehd. &
356, 525, 564, 565, 566, 853, 896, 937, 953 E. H. Wilson 569, 577
Phyllocladus trichomanoides D. Don var. alpinus Picea asperata Mast. var. ponderosa Rehd. &
(Hook. f.) Parl. 390, 391, 523, 566, 644, 853, E. H. Wilson 569, 577
898 Picea asperata Mast. var. retroflexa (Mast.)
Phyllocladus trichomanoides D. Don var. W. C. Cheng 613 1133
trichomanoides 566 Picea aurantiaca Mast. 567, 569, 577, 578, 613,
Picea A. Dietr. 19, 27, 40, 52, 64, 72, 269, 270, 416, 614
417, 431, 451, 459, 463, 474, 479, 484, 511, 567, Picea aurantiaca Mast. var. retroflexa (Mast.)
568, 569, 570, 571, 572, 576, 585, 587, 608, C. T. Kuan & L. J. Zhou 613
610, 652, 687, 740, 789, 963, 1017, 1046, 1049, Picea austromandshurica Silba591
1050 Picea balfouriana Rehd. & E. H. Wilson 597
Picea abies (L.) H. Karst. 27, 62, 68, 499, 567, 571, Picea bicolor (Maxim.) Mayr 573
572, 576, 587, 609, 669, 755, 789, 803 Picea bicolor (Maxim.) Mayr var. acicularis (Maxim.
Picea abies (L.) H. Karst. var. abies569, 572 ex Beissn.) Shiras. 574
Picea abies (L.) H. Karst. var. acuminata (Beck) Picea bicolor (Maxim.) Mayr var. reflexa Shiras.574
Dallim. & A. B. Jacks. 569, 571, 572 Picea brachytyla (Franch.) E. Pritz. 84, 567, 570,
Picea abies (L.) H. Karst. var. alpestris (Brgger) 578, 579, 580, 595, 986
P. A. Schmidt 572 Picea brachytyla (Franch.) E. Pritz. var. ascendens
Picea abies (L.) H. Karst. subsp. obovata (Ledeb.) (Patschke) Silba 580
Hulten607 Picea brachytyla (Franch.) E. Pritz. var. brachytyla
Picea abies (L.) H. Karst. var. obovata (Ledeb.) 77, 579
Lindq. 571, 607 Picea brachytyla (Franch.) E. Pritz. var.
Picea acicularis Maxim. ex Beissn.574 complanata (Mast.) W. C. Cheng ex Rehd.
Picea ajanensis Fisch. ex Carrire591 118, 580, 965
Picea albertiana S. Br. 589 Picea brachytyla (Franch.) E. Pritz. var. pachyclada
Picea albertiana S. Br. var. densata (L. H. Bailey) (Patschke) Silba 579
W. L. Strong & Hills 589 Picea brachytyla (Franch.) E. Pritz. var.
Picea albertiana S. Br. subsp. ogilviei W. L. Strong & rhombisquamea Stapf580
Hills589 Picea breweriana S. Watson 284, 567, 570, 580,
Picea albertiana S. Br. var. porsildii (Raup) 581, 1066
W. L. Strong & Hills 589 Picea chihuahuana Martnez 80, 567, 569, 581,
Picea alcoquiana (Veitch ex Lindl.) Carrire 573, 582, 600, 601, 644
575, 602 Picea complanata Mast.580
Picea alcoquiana (Veitch ex Lindl.) Carrire var. Picea concolor Gordon75
acicularis (Maxim. ex Beissn.) Fitschen 574, Picea crassifolia Kom. 456, 567, 569, 582, 583
575, 594 Picea engelmannii Parry ex Engelm. 101, 117, 415,
Picea alcoquiana (Veitch ex Lindl.) Carrire var. 477, 508, 510, 568, 570, 583, 584, 585, 589, 627,
alcoquiana 573, 575 633, 641, 642, 670, 674, 688, 697, 722, 737, 749,
Picea alcoquiana (Veitch ex Lindl.) Carrire var. 785, 797, 963, 1026
reflexa (Shiras.) Fitschen 574, 575 Picea engelmannii Parry ex Engelm. subsp.
Picea alpestris (Brgger) Stein 572 engelmannii 584
Picea amabilis Douglas ex Loudon63 Picea engelmannii Parry ex Engelm. var. glabra
Picea ascendens Patschke580 Goodman584
Picea engelmannii Parry ex Engelm. subsp. Picea jezoensis (Siebold & Zucc.) Carrire subsp.
Mexicana (Martnez) P. A. Schmidt 129, 130, jezoensis var. komarovii (V. N. Vassil.)
570, 585 W. C. Cheng & L. K. Fu 592
Picea engelmannii Parry ex Engelm. var. mexicana Picea kamtchatkensis Lacass.591
(Martnez) Silba 585 Picea komarovii V. N. Vassil. 592
Picea excelsa (Lam.) Link var. acuminata Beck572 Picea koraiensis Nakai 592, 593, 1023
Picea farreri C. N. Page & Rushforth 567, 570, Picea koraiensis Nakai var. intercedens (Nakai)
585, 586, 1100 Y. L. Chou 593
Picea fennica (Regel) Kom. 567, 571, 586, 587, Picea koraiensis Nakai var. koraiensis 593
607 Picea koraiensis Nakai var. pungsanensis (Uyeki ex
1134 Picea fennica (Regel) Kom. subsp. uralensis (Tepl.) Nakai) Farjon 593
P. A. Schmidt 586 Picea koyamae Shiras. 567, 568, 593, 594, 602
Picea fortunei A. Murray bis 492 Picea likiangensis (Franch.) E. Pritz. 77, 82, 87,
Picea glauca (Moench) Voss 415, 507, 567, 568, 460, 568, 570, 579, 595, 597, 598, 612, 613
584, 587, 591, 598, 599, 615, 658, 674, 1026, 1044, Picea likiangensis (Franch.) E. Pritz. subsp.
1053 balfouriana (Rehd. & E. H. Wilson)
Picea glauca (Moench) Voss subsp. albertiana Rushforth597
(S. Br.) P. A. Schmidt 589 Picea likiangensis (Franch.) E. Pritz. var. balfouriana
Picea glauca (Moench) Voss var. albertiana (S. Br.) (Rehd. & E. H. Wilson) Hillier 597
Sarg.588, 589 Picea likiangensis (Franch.) E. Pritz. var. bhutanica
Picea glauca (Moench) Voss var. densata L. H. Silba596
Bailey589 Picea likiangensis (Franch.) E. Pritz. var. forrestii
Picea glauca (Moench) Voss subsp. engelmannii Silba596
(Engelm.) T. M. C. Taylor 584 Picea likiangensis (Franch.) E. Pritz. var. hirtella
Picea glauca (Moench) Voss var. engelmannii (Rehd. & E. H. Wilson) W. C. Cheng 596
(Engelm.) Boivin 584 Picea likiangensis (Franch.) E. Pritz. var.
Picea glauca (Moench) Voss var. glauca 589 likiangensis 596, 612, 644
Picea glauca (Moench) Voss var. porsildii Raup589 Picea likiangensis (Franch.) E. Pritz. var. linzhiensis
Picea glehnii (F. Schmidt) Mast. 121, 567, 569, 589, W. C. Cheng & L. K. Fu 597
590, 1010 Picea likiangensis (Franch.) E. Pritz. var.
Picea hirtella Rehd. & E. H. Wilson 596 montigena (Mast.) W. C. Cheng 596
Picea hondoensis Mayr592 Picea likiangensis (Franch.) E. Pritz. var. purpurea
Picea intercedens Nakai593 (Mast.) Dallim. & A. B. Jacks. 611
Picea jezoensis (Siebold & Zucc.) Carrire 100, Picea likiangensis (Franch.) E. Pritz. var. rubescens
109, 121, 501, 502, 533, 568, 590, 591, 1010, 1027, Rehd. & E. H. Wilson 118, 126, 578, 597, 614
1048 Picea linzhiensis (W. C. Cheng & L. K. Fu)
Picea jezoensis (Siebold & Zucc.) Carrire var. Rushforth 126, 505, 568, 570, 597
ajanensis (Fisch. ex Carrire) W. C. Cheng & Picea lowiana Gordon75
L. K. Fu 591 Picea lutzii Little568, 598
Picea jezoensis (Siebold & Zucc.) Carrire f. Picea mariana (Mill.) Britton et al. 436, 507, 567,
kamtchatkensis (Lacass.) S. L. Tung & 570, 591, 599, 615, 658, 1024
Y. L. Chou 591 Picea martinezii T. F. Patt. 567, 569, 600, 601
Picea jezoensis (Siebold & Zucc.) Carrire subsp. Picea maximowiczii Regel ex Mast. 567, 568, 601,
hondoensis (Mayr) P. A. Schmidt 86, 128, 602
506, 570, 573, 591, 592 Picea maximowiczii Regel ex Mast. var.
Picea jezoensis (Siebold & Zucc.) Carrire var. maximowiczii 602
hondoensis (Mayr) Rehd. 105, 592 Picea maximowiczii Regel ex Mast. var. senanensis
Picea jezoensis (Siebold & Zucc.) Carrire subsp. Hayashi 602
jezoensis var. jezoensis 591 Picea mexicana Martnez585
Picea meyeri Rehd. & E. H. Wilson 567, 569, 603, Picea shirasawae Hayashi574
623 Picea sitchensis (Bong.) Carrire 19, 63, 91, 117,
Picea meyeri Rehd. & E. H. Wilson var. mongolica 284, 568, 570, 591, 598, 609, 617, 645, 737, 1026,
H. Q. Wu 603 1033, 1052, 1053, 1066
Picea meyeri Rehd. & E. H. Wilson f. pyramidalis Picea smithiana (Wall.) Boiss. 114, 125, 265, 567,
(H. W. Jen & C. G. Bai) L. K. Fu & Nan Li 603 568, 618, 645, 646
Picea meyeri Rehd. & E. H. Wilson var. pyramidalis Picea smithiana (Wall.) Boiss. var. nepalensis Franco
H. W. Jen & C. G. Bai 603 618
Picea montigena Mast.596 Picea spinulosa (Griff.) Beissn. 79, 460, 504, 567,
Picea morinda Link subsp. tianschanica (Rupr.) 570, 597, 598, 619
Berezin616 Picea spinulosa (Griff.) Beissn. var. yatungensis Silba 1135
Picea morrisonicola Hayata 98, 567, 568, 604, 627, 619
636, 638, 644, 695, 986 Picea tianschanica Rupr.616
Picea neoveitchii Mast. 84, 567, 568, 605, 606 Picea torano (Siebold ex K. Koch) Koehne 567,
Picea notabilis (Rehd. & E. H. Wilson) Lacass. 577 568, 620, 621, 622
Picea obovata Ledeb. 96, 109, 124, 464, 498, 501, Picea vulgaris Link var. uralensis Tepl.586
502, 515, 567, 569, 571, 587, 593, 607, 608, 783, Picea watsoniana Mast.622
1101 Picea wilsonii Mast. 567, 568, 603, 612, 622, 623,
Picea obovata Ledeb. var. fennica (Regel) A. Henry 646
586 Picea wilsonii Mast. var. shanxiensis Silba622
Picea omorika (Panci) Purk. 567, 568, 608, 609 Picea wilsonii Mast. var. watsoniana (Mast.) Silba
Picea orientalis (L.) Peterm. 111, 567, 569, 609, 622
610, 645, 818 Pilgerodendron Florin 38, 48, 50, 383, 521, 624
Picea pachyclada Patschke579 Pilgerodendron uviferum (D. Don) Florin 382,
Picea polita (Siebold & Zucc.) Carrire 620 517, 624, 646, 900
Picea ponderosa (Rehd. & E. H. Wilson) Lacass. Pinaceae Spreng. ex F. Rudolphi 18, 19, 20, 24, 27,
577 28, 35, 39, 41, 45, 51, 57, 62, 65, 87, 88, 122, 124,
Picea prostrata Isakov616 130, 258, 259, 261, 274, 488, 491, 497, 504, 514,
Picea pungens Engelm. 568, 570, 610, 611, 627, 549, 567, 586, 593, 599, 612, 626, 705, 741, 954,
629, 775, 777 959, 975, 979, 988, 1001, 1032, 1042
Picea pungsanensis Uyeki ex Nakai593 Pinea Wolf626
Picea purpurea Mast. 84, 118, 568, 570, 611, 612, Pinus L. 15, 20, 26, 27, 29, 35, 39, 40, 51, 52, 57, 76,
623 90, 101, 115, 129, 196, 226, 227, 257, 263, 284, 288,
Picea purpurea Mast. var. balfouriana (Rehd. & 290, 296, 416, 425, 437, 451, 462, 463, 478, 480,
E. H. Wilson) Silba 597 486, 491, 564, 571, 578, 586, 591, 601, 626, 627,
Picea purpurea Mast. var. hirtella (Rehd. & E. H. 628, 630, 631, 632, 633, 634, 635, 653, 658, 673,
Wilson) Silba 596 688, 694, 697, 698, 702, 705, 708, 716, 727, 729,
Picea retroflexa Mast. 567, 569, 578, 613 734, 736, 743, 746, 748, 750, 776, 780, 793, 803,
Picea rubens Sarg. 90, 415, 567, 570, 614, 615, 823, 805, 819, 820, 865, 874, 885, 895, 901, 922, 936,
824, 919, 1044 946, 952, 955, 963, 978, 979, 989, 1025, 1048,
Picea schrenkiana Fisch. & C. A. Mey. 416, 425, 1053, 1065
457, 478, 567, 569, 615 Pinus abies L. 567, 572
Picea schrenkiana Fisch. & C. A. Mey. subsp. Pinus abies L. var. fennica Regel586
schrenkiana 616 Pinus albicaulis Engelm. 19, 443, 508, 627, 635,
Picea schrenkiana Fisch. & C. A. Mey. subsp. 636, 637, 638, 641, 646, 657, 697, 737, 755, 963,
tianschanica (Rupr.) Bykov 616, 645 1066
Picea schrenkiana Fisch. & C. A. Mey. var. Pinus amamiana Koidz. 627, 635, 638, 639
tianschanica (Rupr.) W. C. Cheng & Pinus apulcensis Lindl.768
S. H. Fu 616 Pinus araucana Molina 191, 197
Pinus aristata Engelm. 101, 627, 634, 639, 641, Pinus balfouriana Balf. subsp. longaeva
646, 697, 724, 725 (D. K. Bailey) E. Murray 724
Pinus aristata Engelm. subsp. longaeva Pinus balsamea L. 65
(D. K. Bailey) E. Murray 724 Pinus banksiana Lamb. 19, 65, 415, 436, 507, 599,
Pinus aristata Engelm. var. longaeva (D. K. Bailey) 627, 630, 658, 674, 775, 1024
Little724 Pinus bhutanica Grierson et al. 627, 636, 659
Pinus arizonica Engelm. 300, 627, 633, 641, 642, Pinus bracteata D. Don 68
670, 688, 722, 727, 762, 785, 797 Pinus brutia Ten. 266, 323, 423, 426, 430, 446,
Pinus arizonica Engelm. var. arizonica640, 642 447, 448, 627, 628, 631, 660, 661, 662, 703, 745
Pinus arizonica Engelm. var. cooperi (C. E. Blanco) Pinus brutia Ten. var. brutia 661
1136 Farjon 651 Pinus brutia Ten. var. densifolia F. Yaltirik &
Pinus arizonica Engelm. var. stormiae Martnez M. Boydak 661
651, 701, 774 Pinus brutia Ten. var. eldarica (Medw.) Silba
Pinus armandii Franch. 118, 586, 598, 604, 605, 661
627, 635, 638, 651, 652, 653, 682, 808, 818 Pinus brutia Ten. var. pendulifolia Frankis 662
Pinus armandii Franch. var. amamiana (Koidz.) Pinus brutia Ten. var. pityusa (Steven) Silba 662
Hatus.638 Pinus brutia Ten. var. stankewiczii (Sukaczev)
Pinus armandii Franch. var. armandii646, 652 Frankis662
Pinus armandii Franch. var. dabeshanensis Pinus bungeana Zucc. ex Endl. 29, 489, 627, 635,
(W. C. Cheng & Y. W. Law) Silba 652 647, 662, 663, 699, 784
Pinus armandii Franch. subsp. mastersiana (Hayata) Pinus californiarum D. K. Bailey 733
Businsk653 Pinus californiarum D. K. Bailey subsp. fallax
Pinus armandii Franch. var. mastersiana (Hayata) (Little) D. K. Bailey 733
Hayata98, 653 Pinus canadensis L. 1043
Pinus attenuata Lemmon 313, 314, 320, 322, 406, Pinus canariensis C. Sm. 627, 630, 663, 664, 665
627, 629, 646, 653, 772 Pinus caribaea Morelet 19, 627, 628, 666, 748, 750,
Pinus attenuradiata Stockwell & Righter 654 801, 869
Pinus ayacahuite Ehrenb. ex Schltdl. 19, 92, 94, Pinus caribaea Morelet var. bahamensis (Griseb.)
317, 582, 627, 636, 654, 655, 687, 688, 727, 731, W. H. Barrett & Golfari 401, 667
753, 786, 796 Pinus caribaea Morelet var. caribaea 401, 667,
Pinus ayacahuite Ehrenb. ex Schltdl. var. 801
ayacahuite 655 Pinus caribaea Morelet var. hondurensis (Sncl.)
Pinus ayacahuite Ehrenb. ex Schltdl. var. W. H. Barrett & Golfari 666, 668
novogaliciana Carvajal 785 Pinus catarinae Rob.-Pass. 773
Pinus ayacahuite Ehrenb. ex Schltdl. subsp. Pinus ceciliae Llorens & L. Llorens 702
strobiformis (Engelm.) E. Murray 785 Pinus cembra L. 19, 499, 572, 626, 627, 635, 637,
Pinus ayacahuite Ehrenb. ex Schltdl. var. veitchii 638, 647, 668, 669, 755, 783
(Roezl) Shaw 646, 655, 656 Pinus cembra L. var. pumila Pall. 768
Pinus bahamensis Griseb. 667 Pinus cembra L. subsp. sibirica (Du Tour)
Pinus balfouriana Balf. 627, 634, 647, 656, 657, Krylov782
724, 737 Pinus cembra L. var. sibirica (Du Tour) G. Don
Pinus balfouriana Balf. subsp. aristata (Engelm.) 782
Engelm.639 Pinus cembroides Zucc. 29, 300, 398, 403, 419,
Pinus balfouriana Balf. var. aristata (Engelm.) 420, 429, 445, 450, 473, 627, 634, 669, 670, 686,
Engelm.639 693, 701, 723, 727, 743, 760, 767, 774
Pinus balfouriana Balf. subsp. austrina Bruijn & Pinus cembroides Zucc. subsp. cembroides var.
J. Mastrog. 656, 657 bicolor Little 671
Pinus balfouriana Balf. var. austrina (Bruijn & Pinus cembroides Zucc. subsp. cembroides var.
J. Mastrog.) Silba 656 cembroides 647, 671
Pinus cembroides Zucc. subsp. edulis (Engelm.) Pinus culminicola Andresen & Beaman 19, 627,
E. Murray 689 634, 648, 677, 678
Pinus cembroides Zucc. var. edulis (Engelm.) Pinus culminicola Andresen & Beaman var. discolor
Voss689 (D. K. Bailey & Hawksw.) Silba 671
Pinus cembroides Zucc. var. juarezensis (Lanner) Pinus culminicola Andresen & Beaman var. johannis
Silba770 (Rob.-Pass.) Silba 671
Pinus cembroides Zucc. subsp. lagunae (Rob.- Pinus cupressoides Molina 382
Pass.) D. K. Bailey 671 Pinus dabeshanensis W. C. Cheng & Y. W. Law
Pinus cembroides Zucc. var. lagunae 652
Rob.-Pass.671 Pinus dalatensis Ferr 627, 636, 679, 680, 717, 752,
Pinus cembroides Zucc. subsp. monophylla (Torr. & 806 1137
Frm.) E. Murray 733 Pinus dalatensis Ferr subsp. dalatensis var.
Pinus cembroides Zucc. subsp. orizabensis D. K. bidoupensis Businsk 681
Bailey 648, 672 Pinus dalatensis Ferr subsp. dalatensis var.
Pinus cembroides Zucc. var. orizabensis (D. K. dalatensis 681
Bailey) Silba 672 Pinus dalatensis Ferr subsp. procera Businsk
Pinus cembroides Zucc. var. quadrifolia (Parl. ex 681
Sudw.) Silba 770 Pinus dalmatica Vis. 745
Pinus cembroides Zucc. var. remota Little 773 Pinus dammara Lamb. 148, 155, 156
Pinus chiapensis (Martnez) Andresen 636, 787 Pinus densa (Little & K. W. Dorman) Silba 692
Pinus chihuahuana Engelm. 723 Pinus densa (Little & K. W. Dorman) Silba var.
Pinus cilicica Antoine & Kotschy 74 austrokeysensis Silba 692
Pinus clausa (Chapm. ex Engelm.) Sarg. 627, 630, Pinus densata Mast. 308, 386, 627, 632, 682, 807
672, 673 Pinus densata Mast. var. pygmaea J. R. Xue 808
Pinus clausa (Chapm. ex Engelm.) Sarg. subsp. Pinus densiflora Siebold & Zucc. 86, 97, 286,
immuginata (D. B. Ward) E. Murray 672 461, 506, 602, 622, 627, 632, 682, 683, 684, 794,
Pinus clausa (Chapm. ex Engelm.) Sarg. var. 1055
immuginata D. B. Ward 672, 673 Pinus densiflora Siebold & Zucc. var. densiflora
Pinus clusiana Clemente 313, 443, 510, 747 684
Pinus contorta Douglas ex Loudon 477, 599, 611, Pinus densiflora Siebold & Zucc. var. funebris
627, 630, 637, 673, 674, 697, 711, 737, 963, 1026, (Kom.) T. N. Liou & Q. L. Wang ex Silba683
1066 Pinus densiflora Siebold & Zucc. f. sylvestriformis
Pinus contorta Douglas ex Loudon var. contorta Taken. 683
674 Pinus densiflora Siebold & Zucc. var. sylvestriformis
Pinus contorta Douglas ex Loudon subsp. latifolia (Taken.) Q. L. Wang 683
(Engelm.) Critchf. 675 Pinus densiflora Siebold & Zucc. var. ussuriensis
Pinus contorta Douglas ex Loudon var. latifolia T. N. Liou & Q. L. Wang 684
Engelm.510, 675 Pinus densiflora Siebold & Zucc. var. zhangwuensis
Pinus contorta Douglas ex Loudon subsp. S. J. Zhang & et al. 683, 684
murrayana (Balf.) Engelm. 675 Pinus densi-thunbergii Uyeki 683
Pinus contorta Douglas ex Loudon var. murrayana Pinus densithunbergii Uyeki 627, 683
(Balf.) Engelm. 648, 674, 675, 719 Pinus deodara Lamb.263
Pinus cooperi C. E. Blanco 651 Pinus devoniana Lindl. 92, 627, 633, 648, 685,
Pinus coulteri D. Don 70, 254, 300, 627, 633, 648, 686, 687, 731, 749, 764, 765
675, 711, 712, 781, 961 Pinus discolor D. K. Bailey & Hawksw. 671
Pinus crassicorticea Y. C. Zhong & K. X. Huang Pinus divaricata (Aiton) Dum.-Cours. var. latifolia
729 (Engelm.) Boivin 675
Pinus cubensis Griseb. 627, 628, 676, 677 Pinus donnell-smithii Mast. 703
Pinus douglasiana Martnez 627, 633, 686, 687, Pinus funebris Kom. 683, 684
710, 722, 727, 732, 749, 765 Pinus gerardiana Wall. ex D. Don 29, 428, 627,
Pinus douglasiana Martnez var. martinezii 635, 698, 699, 784
(E. Larsen) Silba 687 Pinus glabra Walter 627, 628, 629, 699, 1040
Pinus douglasiana Martnez var. maximinoi Pinus glauca Moench 589
(H. E. Moore) Silba 731 Pinus gordoniana Hartw. ex Gordon736
Pinus douglasii Sabine ex D. Don 959 Pinus grandis Douglas ex D. Don 90
Pinus dumosa D. Don 1048 Pinus greggii Engelm. ex Parl. 627, 629, 700, 753
Pinus durangensis Martnez 80, 627, 633, 648, Pinus greggii Engelm. ex Parl. var. australis
687, 688, 722, 785, 797 Donahue & Lopez 701
1138 Pinus echinata Mill. 486, 627, 629, 688, 689, 700, Pinus greggii Engelm. ex Parl. var. greggii 701
750, 777, 804 Pinus hakkodensis Makino 627, 702
Pinus edulis Engelm. 29, 300, 440, 445, 477, 626, Pinus halepensis Mill. 29, 446, 448, 627, 631, 661,
627, 634, 689, 690 702, 703, 745, 761
Pinus edulis Engelm. var. fallax Little 733 Pinus halepensis Mill. subsp. brutia (Ten.) Holmboe
Pinus eldarica Medw. 661 661
Pinus elliottii Engelm. 28, 486, 627, 630, 691, 700, Pinus halepensis Mill. var. brutia (Ten.) A. Henry
750, 782 661
Pinus elliottii Engelm. subsp. densa (Little & Pinus halepensis Mill. var. ceciliae (Llorens &
K. W. Dorman) E. Murray 692 L. Llorens) Rosell & et al. 702
Pinus elliottii Engelm. var. densa Little & Pinus hamata (Steven) Sosn. 790
K. W. Dorman 692 Pinus hartwegii Lindl. 92, 120, 317, 442, 480, 627,
Pinus elliottii Engelm. var. elliottii 692 633, 649, 678, 686, 703, 704, 705, 785
Pinus engelmannii Carrire 627, 642, 648, 692, Pinus hartwegii Lindl. var. rudis (Endl.) Silba 703
693 Pinus hayatana Businsk 738
Pinus engelmannii Carrire var. blancoi (Martnez) Pinus heldreichii H. Christ 627, 628, 630, 631, 649,
Martnez692 705, 706, 745
Pinus eremitana Businsk 694 Pinus heldreichii H. Christ subsp. leucodermis
Pinus escarena Risso 757 (Antoine) E. Murray 705
Pinus estevezii (Martnez) J. P. Perry 767 Pinus heldreichii H. Christ var. leucodermis
Pinus fallax (Little) Businsk 733 (Antoine) Markgr. ex Fitschen705
Pinus fenzeliana Hand.-Mazz. 180, 260, 537, 551, Pinus henryi Mast. 706, 707
627, 636, 638, 652, 694, 752, 806, 1008 Pinus herrerae Martnez 707
Pinus fenzeliana Hand.-Mazz. var. annamiensis Pinus hingganensis H. J. Zhang 782
Silba694 Pinus holfordiana A. B. Jacks. 655
Pinus fenzeliana Hand.-Mazz. var. dabeshanensis Pinus hondurensis Sncl. 668
(W. C. Cheng & Y. W. Law) L. K. Fu & Nan Li Pinus hwangshanensis W.Y. Hsia 627, 632, 708,
652 709, 794
Pinus flexilis E. James 101, 415, 477, 508, 627, 635, Pinus inops Aiton var. clausa Chapm. ex Engelm.
655, 695, 696, 697, 698, 724, 737, 786, 963 672
Pinus flexilis E. James var. flexilis 697 Pinus insignis Douglas ex Loudon var. binata
Pinus flexilis E. James var. macrocarpa Engelm. Engelm.773
697 Pinus insularis Endl. 713, 807, 824, 826
Pinus flexilis E. James subsp. reflexa (Engelm.) Pinus insularis Endl. var. khasyana (Griff.)
E. Murray 697 Silba713
Pinus flexilis E. James var. reflexa Engelm. 697 Pinus insularis Endl. var. langbianensis (A. Chev.)
Pinus formosana Hayata 738 Silba713
Pinus fragilissima Businsk 794, 795 Pinus insularis Endl. var. tenuifolia (W. C. Cheng &
Pinus fraseri Pursh 89 Y. W. Law) Silba 808
Pinus insularis Endl. var. yunnanensis (Franch.) Pinus latteri Mason 627, 631, 649, 719, 720, 732,
Silba808 733
Pinus jaliscana Prez de la Rosa 627, 629, 709, Pinus lawsonii Roezl ex Gordon 627, 630, 687,
710 721, 722, 765
Pinus jeffreyi Balf. 254, 300, 304, 443, 627, 633, Pinus lawsonii Roezl ex Gordon var. gracilis
710, 711, 712, 719, 734, 737, 763, 771, 961, 982 Debreczy & Rcz 721
Pinus jeffreyi Balf. var. baja-californica Silba 710 Pinus leiophylla Schiede ex Schltdl. &
Pinus johannis Rob.-Pass. 671 Cham. 300, 419, 627, 628, 687, 688, 693, 721,
Pinus juarezensis Lanner 770 722, 723, 726, 749, 797
Pinus kaempferi Lamb. 505 Pinus leiophylla Schiede ex Schltdl. & Cham. subsp.
Pinus kesiya Royle ex Gordon 19, 28, 627, 628, chihuahuana (Engelm.) E. Murray 723 1139
632, 681, 712, 713, 733, 807, 808 Pinus leiophylla Schiede ex Schltdl. & Cham. var.
Pinus kesiya Royle ex Gordon subsp. insularis chihuahuana (Engelm.) Shaw 80, 670, 722,
(Endl.) D. Z. Li 713 723, 730, 785
Pinus kesiya Royle ex Gordon var. kesiya 713 Pinus leiophylla Schiede ex Schltdl. & Cham. var.
Pinus kesiya Royle ex Gordon var. langbianensis Leiophylla 722, 723
(A. Chev.) Gaussen ex N.-S. Bui 713, 807 Pinus leucodermis Antoine 705, 706
Pinus kesiya Royle ex Gordon subsp. yunnanensis Pinus longaeva D. K. Bailey 19, 627, 634, 641, 649,
(Franch.) Businsk 808 657, 697, 724, 725, 734, 743
Pinus kochiana Klotzsch ex K. Koch 790 Pinus longipedunculata (Loock ex Martnez)
Pinus koraiensis Siebold & Zucc. 29, 96, 100, 109, Businsk754
461, 464, 533, 567, 569, 627, 635, 713, 714, 1010, Pinus luchuensis Mayr 627, 632, 709, 725, 794
1023 Pinus luchuensis Mayr subsp. hwangshanensis
Pinus krempfii Lecomte 626, 627, 634, 679, 715, (W. Y. Hsia) D. Z. Li 708
716, 717 Pinus luchuensis Mayr var. hwangshanensis (W. Y.
Pinus krempfii Lecomte var. poilanei Lecomte Hsia) C. L. Wu 708
716 Pinus lumholtzii B.L. Rob & Fernald 627, 628,
Pinus kwangtungensis Chun & Tsiang 175, 258, 693, 726, 727, 764, 785
260, 277, 537, 551, 694, 752, 806, 1008 Pinus lumholtzii B. L. Rob & Fernald var.
Pinus kwangtungensis Chun & Tsiang var. variifolia microphylla Carvajal 723
Nan Li & Y. C. Zhong 694 Pinus luzmariae Prez de la Rosa 627, 629, 727
Pinus lagunae (Rob.-Pass.) Passini 671 Pinus macrophylla Engelm. var. blancoi Martnez
Pinus lambertiana Douglas 19, 254, 300, 322, 581, 692
627, 636, 711, 717, 718, 737, 982, 1026 Pinus macvaughii Carvajal 709
Pinus lambertiana Douglas var. martirensis Silba Pinus maestrensis Bisse 676
717 Pinus martinezii E. Larsen 627, 687
Pinus lanceolata Lamb. 294, 296 Pinus massoniana Lamb. 28, 297, 386, 489, 551,
Pinus langbianensis A. Chev. 713 627, 632, 684, 707, 727, 728
Pinus laricina Du Roi 506 Pinus massoniana Lamb. var. hainanensis
Pinus laricio Poir. 746 W. C. Cheng & L. K. Fu 729
Pinus laricio Poir. subsp. calabrica (Loudon) Cesca Pinus massoniana Lamb. var. henryi (Mast.)
& Peruzzi 746 C. L. Wu 706
Pinus laricio Poir. var. angustisquama Willk. 747 Pinus massoniana Lamb. var. massoniana 729
Pinus laricio Poir. var. calabrica Loudon 746 Pinus massoniana Lamb. var. shaxianensis
Pinus laricio Poir. var. caramanica Loudon 746 D. X. Zhou 729
Pinus laricio Poir. var. latisquama Willk. 747 Pinus massoniana Lamb. var. wulingensis C. J. Qi &
Pinus larix L. var. russica Endl. 506, 514 Q. Z. Lin 706
Pinus lasiocarpa Hook. 101 Pinus mastersiana Hayata 653
Pinus maximartinezii Rzed. 19, 627, 635, 649, 729, Pinus mugo Turra subsp. uncinata (Ramond ex DC)
730 Domin 802
Pinus maximinoi H. E. Moore 317, 627, 633, 686, Pinus mukdensis Uyeki ex Nakai792
710, 731, 749, 753, 765, 796 Pinus muricata D. Don 313, 627, 628, 650, 654,
Pinus merkusii Jungh. & de Vriese 627, 631, 720, 741
732, 733 Pinus muricata D. Don var. borealis Axelrod ex
Pinus merkusii Jungh. & de Vriese subsp. latteri Farjon741
(Mason) D. Z. Li 719 Pinus muricata D. Don var. cedrosensis J. T. Howell
Pinus merkusii Jungh. & de Vriese var. latteri 773
(Mason) Silba 719 Pinus muricata D. Don var. stantonii Axelrod ex
1140 Pinus merkusii Jungh. & de Vriese subsp. ustulata Farjon741
Businsk732 Pinus murrayana Balf. 675
Pinus mertensiana Bong. 1042, 1053 Pinus neilreichiana Reichardt 627, 742
Pinus mesogeensis Fieschi & Gaussen 756, 757 Pinus nelsonii Shaw 450, 627, 634, 650, 670, 742,
Pinus michoacana Martnez 685 743
Pinus monophylla Torr. & Frm. 29, 300, 445, 627, Pinus nigra J. F. Arnold 71, 74, 266, 423, 426, 430,
631, 649, 650, 733, 734, 771 446, 572, 609, 627, 630, 632, 684, 705, 742, 744,
Pinus monophylla Torr. & Frm. var. fallax (Little) 746, 798
Silba733 Pinus nigra J. F. Arnold var. angustisquama (Willk.)
Pinus monophylla Torr. & Frm. var. californiarum Laguna Lumbreras 747
(D. K. Bailey) Silba 733 Pinus nigra J. F. Arnold subsp. calabrica (Loudon)
Pinus montezumae Lamb. 92, 94, 120, 317, 398, E. Murray 746
627, 633, 686, 688, 701, 703, 704, 721, 722, 734, Pinus nigra J. F. Arnold var. calabrica (Loudon) C.K.
735, 797 Schneid.746
Pinus montezumae Lamb. var. gordoniana (Hartw. Pinus nigra J. F. Arnold subsp. caramanica
ex Gordon) Silba 736 (Loudon) Businsk 746
Pinus montezumae Lamb. var. mezambrana Pinus nigra J. F. Arnold var. caramanica (Loudon)
Carvajal736 Rehd.746
Pinus montezumae Lamb. var. montezumae 736 Pinus nigra J. F. Arnold subsp. clusiana (Clemente)
Pinus montezumae Lamb. var. rudis (Endl.) Shaw Rivas-Martnez747
703 Pinus nigra J. F. Arnold var. columnaris-pendula
Pinus monticola Douglas ex D. Don 254, 443, 510, Boydak746
581, 627, 636, 655, 711, 719, 736, 737, 763, 805, Pinus nigra J. F. Arnold subsp. croatica Lovric 745
1026, 1066 Pinus nigra J. F. Arnold subsp. dalmatica (Vis.)
Pinus morrisonicola Hayata 627, 636, 695, 738, Franco 745
752 Pinus nigra J. F. Arnold var. dalmatica (Vis.)
Pinus mugo Turra 416, 627, 631, 678, 739, 740, 745, Businsk745
776, 802 Pinus nigra J. F. Arnold subsp. laricio (Poir.)
Pinus mugo Turra subsp. mugo 740 Maire 650, 746
Pinus mugo Turra subsp. rostrata (Antoine) Pinus nigra J. F. Arnold var. latisquama (Willk.)
E. Murray 802 Laguna Lumbreras 747
Pinus mugo Turra var. rostrata (Antoine) Hoopes Pinus nigra J. F. Arnold subsp. nigra 705, 742, 745
802 Pinus nigra J. F. Arnold subsp. pallasiana (Lamb.)
Pinus mugo Turra subsp. rotundata (Link) Janch. Holmboe447, 746
& H. Neumayer 739, 740, 802 Pinus nigra J. F. Arnold subsp. pallasiana (Lamb.)
Pinus mugo Turra subsp. rotundata (Link) Janch. & Holmboe var. fastigiata Businsk 746
H. Neumayer var. pseudopumilio (Willk.) Pinus nigra J. F. Arnold subsp. pallasiana (Lamb.)
P. Schmidt 740 Holmboe var. pallasiana 746
Pinus nigra J. F. Arnold var. pyramidata A. Acatay Pinus parviflora Siebold & Zucc. var. fenzeliana
(nom. inval.)747 (Hand.-Mazz.) C.L. Wu694
Pinus nigra J. F. Arnold subsp. salzmannii (Dunal) Pinus parviflora Siebold & Zucc. var. morrisonicola
Franco 650, 747, 757 (Hayata) C. L. Wu 738
Pinus nigra J. F. Arnold var. salzmannii (Dunal) Pinus parviflora Siebold & Zucc. var. parviflora
Laguna Lumbreras 747 752
Pinus nigra J. F. Arnold var. yaltirikiana Pinus parviflora Siebold & Zucc. subsp. pentaphylla
C.U. Alptekin746 (Mayr) Businsk 752
Pinus nigra J. F. Arnold var. fastigiata Pinus parviflora Siebold & Zucc. var. pentaphylla
Businsk 746 (Mayr) A. Henry 702, 752
Pinus nordmanniana Steven 111 Pinus patula Schiede ex Schltdl. & Cham. 95, 317, 1141
Pinus novaemexicana P. Landry 697 627, 628, 701, 722, 731, 753, 765, 797, 1061
Pinus nubicola J. P. Perry 768 Pinus patula Schiede ex Schltdl. & Cham. var.
Pinus oaxacana Mirov 768 jaliscana (Prez de la Rosa) Silba 709
Pinus oaxacana Mirov var. diversiformis Debreczy Pinus patula Schiede ex Schltdl. & Cham. var.
& Rcz 768 longipedunculata Loock ex Martnez 754
Pinus occidentalis Sw. 401, 627, 629, 747, 748 Pinus patula Schiede ex Schltdl. & Cham. var.
Pinus occidentalis Sw. var. baorucoensis Silba 747 patula 650, 754
Pinus occidentalis Sw. var. cubensis (Griseb.) Silba Pinus patula Schiede ex Schltdl. & Cham. subsp.
676 tecunumanii (Eguiluz & J. P. Perry) Styles 795
Pinus occidentalis Sw. var. maestrensis (Bisse) Silba Pinus pentaphylla Mayr 752
676 Pinus pentaphylla Mayr var. hakkodensis (Makino)
Pinus omorika Pani 608 Kusaka702
Pinus oocarpa Schiede ex Schltdl. 433, 627, 629, Pinus peuce Griseb. 627, 636, 745, 754, 755
668, 686, 687, 688, 710, 721, 722, 727, 731, 748, Pinus peuce var. vermiculata Christ 755
749, 764, 765, 779, 796, 797, 869 Pinus picea L. 57
Pinus oocarpa Schiede ex Schltdl. var. macvaughii Pinus pinaster Aiton 28, 115, 627, 628, 630, 756,
(Carvajal) Silba 709 757, 758
Pinus oocarpa Schiede ex Schltdl. var. manzanoi Pinus pinaster Aiton subsp. acutisquama (Boiss.)
Martnez748 Rivas-Martnez757
Pinus oocarpa Schiede ex Schltdl. var. ochoterenae Pinus pinaster Aiton var. acutisquama Boiss. 757
Martnez795 Pinus pinaster Aiton subsp. atlantica Villar 756, 757
Pinus oocarpa Schiede ex Schltdl. f. trifoliata Pinus pinaster Aiton subsp. escarena (Risso) K.
Martnez727 Richt. 757
Pinus oocarpa Schiede ex Schltdl. var. trifoliata Pinus pinaster Aiton var. mesogeensis (Fieschi &
Martnez727 Gaussen) Silba 757
Pinus orientalis L. 609 Pinus pinaster Aiton subsp. pinaster 757, 809
Pinus orizabensis (D. K. Bailey) D. K. Bailey & Pinus pinaster Aiton subsp. renoui (Villar)
Hawksw.672 Maire756, 758
Pinus orthophylla Businsk 694 Pinus pinaster Aiton var. renoui Villar 758
Pinus pallasiana Lamb. 746 Pinus pinceana Gordon 450, 627, 635, 670, 743,
Pinus pallasiana Lamb. subsp. caramanica 758, 759
(Loudon) Chrtek & B. Slavik 746 Pinus pindrow Royle ex D. Don 114
Pinus palustris Mill. 19, 28, 627, 629, 686, 750, 751, Pinus pinea L. 29, 446, 627, 631, 626, 745, 760, 761,
777, 1013 809
Pinus parviflora Siebold & Zucc. 86, 128, 286, Pinus pityusa Steven 662
573, 627, 636, 738, 751, 752, 806, 978, 1027, 1030, Pinus pityusa Steven var. stankewiczii
1048, 1055 Sukaczev662
Pinus ponderosa Douglas ex C. Lawson 70, 254, Pinus radiata D. Don 25, 236, 313, 315, 321, 627,
300, 304, 313, 314, 320, 420, 443, 477, 510, 581, 629, 654, 701, 742, 771, 772, 923
627, 628, 632, 633, 642, 690, 711, 712, 719, 721, Pinus radiata D. Don subsp. binata (Engelm.)
734, 737, 761, 762, 763, 961, 963, 982, 1026 E. Murray 773
Pinus ponderosa Douglas ex C. Lawson subsp. Pinus radiata D. Don var. binata (Engelm.)
arizonica (Engelm.) E. Murray 642 Lemmon315, 773
Pinus ponderosa Douglas ex C. Lawson var. Pinus radiata D. Don var. cedrosensis (J. T. Howell)
arizonica (Engelm.) Shaw 642 Silba773
Pinus ponderosa Douglas ex C. Lawson subsp. Pinus radiata D. Don var. radiata 773
coulteri (D. Don) E. Murray 675 Pinus remota (Little) D. K. Bailey & Hawksw. 399,
1142 Pinus ponderosa Douglas ex C. Lawson var. 627, 634, 743, 773, 774
ponderosa 763, 809 Pinus reflexa (Engelm.) Engelm. 697
Pinus ponderosa Douglas ex C. Lawson subsp. Pinus religiosa Kunth 119
scopulorum (Engelm.) E. Murray 763 Pinus renoui (Villar) Gaussen 758
Pinus ponderosa Douglas ex C. Lawson var. Pinus resinosa Aiton 627, 632, 658, 774, 775, 1044
scopulorum Engelm. 763 Pinus rhaetica Brgger 627, 776
Pinus ponderosa Douglas ex C. Lawson var. Pinus rigida Mill. 627, 629, 770, 775, 776, 781, 782,
stormiae (Martnez) Silba 651 804
Pinus ponderosa Douglas ex C. Lawson subsp. Pinus rotundata Link 740
washoensis (Mason & Stockw.) E. Murray 763 Pinus roxburghii Sarg. 627, 630, 660, 777, 778,
Pinus praetermissa Styles & McVaugh 627, 629, 779, 805
764 Pinus rudis Endl. 703
Pinus pringlei Shaw 627, 630, 721, 722, 764, 765 Pinus rzedowskii Madrigal & M. Caball. 627, 635,
Pinus prokoraiensis Y. T. Zhao 713 779, 810, 1095
Pinus pseudostrobus Lindl. 92, 94, 95, 317, 582, Pinus sabiniana Douglas ex D. Don 300, 320, 322,
627, 633, 670, 686, 687, 693, 701, 721, 722, 731, 406, 627, 633, 780, 781
732, 753, 764, 765, 766, 767, 779, 796 Pinus salzmannii Dunal 747
Pinus pseudostrobus Lindl. subsp. apulcensis (Lindl.) P. schwerinii Fitschen 805
Stead768 Pinus scopulorum (Engelm.) Lemmon 763
Pinus pseudostrobus Lindl. var. apulcensis (Lindl.) Pinus serotina Michx. 627, 629, 691, 781, 782
Martnez768 Pinus sibirica Du Tour 109, 124, 457, 515, 593, 627,
Pinus pseudostrobus Lindl. var. apulcensis (Lindl.) 635, 782, 783, 1023
Shaw 768 Pinus sibirica Du Tour var. hingganensis
Pinus pseudostrobus Lindl. var. estevezii Martnez (H. J. Zhang) Silba 782
767 Pinus sitchensis Bong. 617
Pinus pseudostrobus Lindl. var. laubenfelsii Silba Pinus smithiana Wall. 618
768 Pinus sosnowskyi Nakai 790
Pinus pseudostrobus Lindl. var. oaxacana (Mirov) Pinus spectabilis D. Don 124
S. G. Harrison 768 Pinus squamata X. W. Li 627, 635, 783
Pinus pseudostrobus Lindl. var. Pinus strobiformis Engelm. 80, 81, 582, 627, 636,
pseudostrobus 767 642, 656, 698, 785, 786
Pinus pumila (Pall.) Regel 19, 105, 109, 121, 128, Pinus strobiformis Englem. subsp. veitchii (Roezl)
464, 501, 502, 533, 591, 627, 635, 678, 702, 768, Frankis656
1023 Pinus strobiformis Engelm. var. carvajalii Silba 785
Pinus pungens Lamb. 568, 570, 627, 629, 769, 770, Pinus strobiformis Engelm. var. potosiensis
775, 777, 809 Silba697
Pinus quadrifolia Parl. ex Sudw. 627, 635, 734, Pinus strobus L. 64, 317, 582, 615, 626, 627, 636,
770, 771, 809 653, 655, 737, 765, 775, 785, 786, 1024, 1044
Pinus strobus L. subsp. chiapensis (Martnez) Pinus taiwanensis Hayata 295, 627, 632, 709, 793,
E. Murray 787 794
Pinus strobus L. var. chiapensis Martnez 737, Pinus taiwanensis Hayata var. damingshanensis
787, 796, 1014 W. C. Cheng & L. K. Fu 708, 709
Pinus strobus L. subsp. monticola (Douglas ex Pinus taiwanensis Hayata var. fragilissima
D. Don) E. Murray 736 (Businsk) Farjon 795
Pinus strobus L. var. monticola (Douglas ex D. Don) Pinus taiwanensis Hayata var. taiwanensis 794,
Nuttall736 810
Pinus strobus L. var. strobus 787 Pinus tecunumanii Eguiluz & J. P. Perry 627, 629,
Pinus stylesii Frankis ex Businsk697 668, 731, 749, 795, 796
Pinus sylvestris L. 19, 27, 62, 111, 414, 498, 500, 501, Pinus tenuifolia Benth 731 1143
515, 571, 587, 608, 626, 627, 682, 684, 742, 744, Pinus teocote Schiede ex Schltdl. & Cham. 29, 95,
745, 769, 776, 783, 788, 789, 803, 825, 883, 1094, 419, 627, 688, 701, 721, 722, 727, 796
1109 Pinus teocote Schiede ex Schltdl. & Cham. var.
Pinus sylvestris L. subsp. hamata (Steven) Fomin herrerae (Martnez) Silba 707
790 Pinus thunbergiana Franco 797
Pinus sylvestris L. subsp. kochiana (Klotzsch ex Pinus thunbergii Parl. 627, 631, 683, 684, 797,
K. Koch) Eliin 790 798
Pinus sylvestris L. var. manguiensis S. Y. Li & Pinus torreyana Parry ex Carrire 627, 633, 798,
Adair790 799
Pinus sylvestris L. var. scotica Beissn. 789 Pinus torreyana Parry ex Carrire subsp. insularis
Pinus sylvestris L. var. sylvestriformis (Taken.) J. R. Haller 800
W. C. Cheng & C. D. Chu 683 Pinus torreyana Parry ex Carrire var. insularis
Pinus sylvestris L. var. hamata Steven 790 (J. R. Haller) Silba 800
Pinus sylvestris L. var. mongolica Litv. 790 Pinus torreyana Parry ex Carrire subsp.
Pinus sylvestris L. var. sylvestris 788, 789, 810 Torreyana 799, 800
Pinus tabuliformis Carrire 297, 627, 632, 663, Pinus tropicalis Morelet 401, 627, 632, 801, 802
682, 707, 790, 791, 807, 808, 818 Pinus uliginosa Neuman ex Wimm.740
Pinus tabuliformis Carrire var. brevifolia S. Y. Wang Pinus uncinata Ramond ex DC. 627, 631, 740,
& C. L. Chang 791 802, 803
Pinus tabuliformis Carrire var. densata (Mast.) Pinus uncinata Ramond ex DC. var. ancestralis
Rehd.682 Businsk802
Pinus tabuliformis Carrire subsp. henryi (Mast.) Pinus uncinata Ramond ex DC. var. pseudopumilio
Businsk706 Willk.740
Pinus tabuliformis Carrire var. henryi (Mast.) Pinus uncinata Ramond ex DC. var. rostrata
C. T. Kuan 706 Antoine802
Pinus tabuliformis Carrire subsp. mukdensis (Uyeki Pinus uncinata Ramond ex DC. subsp. uliginosa
ex Nakai) Businsk 792 (Neumann ex Wimm.) Businsk 740
Pinus tabuliformis Carrire var. mukdensis (Uyeki Pinus uyematsui Hayata 738
ex Nakai) Uyeki 792 Pinus veitchii Roezl 656
Pinus tabuliformis Carrire var. pygmaea (J. R. Xue) Pinus virginiana Mill. 486, 627, 630, 770, 775,
Silba808 777, 803, 804, 810
Pinus tabuliformis Carrire var. tabuliformis 791 Pinus wallichiana A. B. Jacks. 114, 125, 265, 428,
Pinus tabuliformis Carrire var. umbraculifera 457, 460, 478, 504, 579, 618, 620, 627, 636, 655,
T. N. Liou & Q. L. Wang 792 659, 660, 681, 778, 804, 810
Pinus tabuliformis Carrire var. yunnanensis Dallim. Pinus wallichiana A. B. Jacks. subsp. bhutanica
& A.B. Jacks.808 (Grierson et al.) Businsk 659
Pinus taeda L. 627, 629, 689, 691, 700, 750, 781, Pinus wallichiana A. B. Jacks. var. manangensis
792, 804, 1040 H. Ohba & M. Suzuki 805
Pinus wallichiana A. B. Jacks. var. parva Podocarpus alpinus R. Br. ex Hook. f. var.
K. C. Sahni 805 arborescens Brongn. & Gris 866
Pinus wallichiana A. B. Jacks. var. wallichiana Podocarpus alpinus R. Br. ex Hook. f. var.
805 caespitosus Pancher ex Brongn. & Gris 866
Pinus wangii Hu & W. C. Cheng 627, 636, 694, Podocarpus alpinus R. Br. ex Hook. f. var. lawrencei
752, 806, 807 (Hook. f.) Hook. f. 878
Pinus wangii Hu & W. C. Cheng subsp. kwangtungensis Podocarpus amarus Blume 983
(Chun & Tsiang) Businsk 694 Podocarpus andinus Poepp. ex Endl. 943, 944
Pinus wangii Hu & W. C. Cheng var. Podocarpus angustifolius Griseb. 832, 833, 858
kwangtungensis (Chun & Tsiang) Silba 694 Podocarpus angustifolius Griseb. var. aristulatus
1144 Pinus wangii Hu & W. C. Cheng subsp. variifolia (Parl.) Staszk. 832
(Nan Li & Y. C. Zhong) Businsk 694 Podocarpus angustifolius Griseb. subsp. buchii
Pinus washoensis Mason & Stockw. 763 (Urb.) Staszk. 843
Pinus yecorensis Debreczy & Rcz 768 Podocarpus angustifolius Griseb. subsp. buchii
Pinus yecorensis Debreczy & Rcz var. sinaloensis (Urb.) Staszk. var. latifolius (Florin)
Debreczy & Rcz 768 Staszk.843
Pinus yunnanensis Franch. 627, 632, 682, 784, Podocarpus angustifolius Griseb. var. leonii
807, 808 (Carabia) Staszk. 832
Pinus yunnanensis Franch. var. pygmaea Podocarpus angustifolius Griseb. var. leonii
(J. R. Xue) J. R. Xue 808 (Carabia) Staszk. f. victorinianus (Carabia)
Pinus yunnanensis Franch. var. tenuifolia W. C. Staszk.832
Cheng & Y. W. Law 807, 808 Podocarpus angustifolius Parl. 904
Pinus yunnanensis Franch. var. yunnanensis 807, Podocarpus annamiensis N. E. Gray 895, 897
808 Podocarpus annamiensis N.E. Gray var. hainanensis
Platycladus Spach 38, 48, 50, 532, 817, 818 Gaussen897
Platycladus chengii (Bordres & Gaussen) Podocarpus aracensis de Laub. & Silba 834
A. V. Bobrov 817 Podocarpus archboldii N. E. Gray 835
Platycladus orientalis (L.) Franco 409, 810, 817, Podocarpus archboldii N. E. Gray var. crassiramosus
818, 1098, 1100, 1109 N. E. Gray 912
Platycyparis A. V. Bobrov & Melikyan 298 Podocarpus argotaenia Hance 174, 175
Platycyparis funebris (Endl.) A. V. Bobrov & Podocarpus aristulatus Parl. 832, 833
Melikyan 298, 311 Podocarpus aristulatus Parl. var. buchii (Urb.) Silba
Podocarpaceae Endl. 13, 14, 17, 18, 20, 21, 24, 26, 843
28, 35, 41, 45, 50, 51, 53, 133, 140, 210, 214, 258, Podocarpus aspleniifolius Labill. 560, 561
327, 347, 364, 372, 373, 375, 379, 389, 495, 516, Podocarpus atjehensis (Wasscher) de Laub. ex Silba
528, 529, 534, 536, 555, 557, 563, 819, 877, 879, 912, 836
943, 947, 951, 953, 967, 968, 975, 983, 1078, 1071, Podocarpus ballivianensis Silba 901
1087, 1093, 1094, 1096, 1098, 1100, 1102 Podocarpus barretoi de Laub. & Silba 839
Podocarpus LHr. ex Pers. 15, 20, 24, 28, 41, 53, Podocarpus beecherae de Laub. 898, 899
54, 140, 143, 144, 155, 208, 228, 327, 329, 330, 333, Podocarpus biformis Hook. 390
334, 347, 358, 491, 522, 542, 553, 555, 563, 819, Podocarpus borneensis de Laub. 837
820, 821, 822, 823, 825, 826, 827, 828, 837, 844, Podocarpus bracteatus Blume 838
854, 869, 870, 873, 889, 896, 913, 929, 968, 973, Podocarpus brasiliensis de Laub. 839
983 Podocarpus brassii Pilg. 840, 842
Podocarpus acuminatus de Laub. 830 Podocarpus brassii Pilg. var. brassii 811, 841
Podocarpus acutifolius Kirk 811, 829, 830, 831 Podocarpus brassii Pilg. var. humilis de Laub.
Podocarpus affinis Seem. 831, 832 841
Podocarpus allenii Standl. 868 Podocarpus brevifolius (Stapf) Foxw. 336, 360,
Podocarpus alpinus R. Br. ex Hook. f. 878 811, 841, 843, 1068
Podocarpus brownii C. E. Bertrand 856 Podocarpus drouynianus F. Muell. 825, 856, 857,
Podocarpus buchholzii de Laub. 916 892
Podocarpus buchholzii de Laub. var. neblinensis Podocarpus ekmanii Urb. 826, 857, 858
Silba916 Podocarpus elatus R. Br. ex Endl.238, 812, 825,
Podocarpus buchii Urb. 843, 844 829, 831, 858, 859, 860
Podocarpus buchii Urb. var. latifolius Florin 843 Podocarpus elongatus (Aiton) LHerit. ex
Podocarpus capuronii de Laub. 844, 845 Pers. 819, 820, 829, 860, 861
Podocarpus capuronii de Laub. var. woltzii Podocarpus ensiculus Melville 870
(Gaussen) Silba 844 Podocarpus epiphyticus de Laub. & Silba 896, 897
Podocarpus cardenasii J. T. Buchholz & N. E. Gray Podocarpus expansus (de Laub.) Whitmore 332
865 Podocarpus falcatus (Thunb.) Endl. 142 1145
Podocarpus celatus de Laub. 828, 845, 846 Podocarpus falciformis Parl. 375, 376
Podocarpus celebicus Warb. 908 Podocarpus fasciculus de Laub. 821, 807, 862, 863,
Podocarpus chinensis Wall. ex J. Forbes 886 932
Podocarpus chinensis Wall. ex J. Forbes var. wardii Podocarpus ferrugineus G. Benn. ex D. Don 946
de Laub. & Silba 886 Podocarpus ferruginoides R. H. Compton 947
Podocarpus chingianus (N. E. Gray) S. Y. Hu 811, Podocarpus filicifolius N. E. Gray 972
821, 846, 847, 885 Podocarpus fleuryi Hickel 537
Podocarpus cinctus Pilg. 328 Podocarpus formosensis Dummer 540
Podocarpus colliculatus (N. E. Gray) de Laub 933 Podocarpus formosensis Dummer var. koshuensis
Podocarpus compactus Wasscher 329 (Kaneh.) Merr. & Yamam. 540
Podocarpus comptonii J. T. Buchholz 967 Podocarpus gaussenii Woltz 142, 143
Podocarpus confertus de Laub. 823, 847 Podocarpus gibbsiae N. E. Gray 359, 823, 863
Podocarpus coriaceus Rich. 826, 848, 939 Podocarpus glaucus Foxw. 564, 822, 824, 864, 873
Podocarpus costalis C. Presl 811, 821, 829, 849, Podocarpus globulus de Laub. 823, 865
914 Podocarpus glomeratus D. Don 827, 865, 866
Podocarpus costalis C. Presl var. taiwanensis Podocarpus gnidioides Carrire 825, 866, 867
Gaussen849 Podocarpus gnidioides Carrire var. caespitosus
Podocarpus costaricensis de Laub. ex Silba826, Pancher ex Carrire866
850, 851 Podocarpus gracilimus Stapf 142, 143
Podocarpus crassigemma de Laub. 824, 851, 852 Podocarpus gracilior Pilg. 143
Podocarpus cumingii Parl. 327 Podocarpus grayae de Laub. 812, 867, 868
Podocarpus cunninghamii Colenso 332, 392, 523, Podocarpus guatemalensis Standl. 827, 828, 868,
525, 566, 811, 826, 831, 852, 853, 937 869
Podocarpus cupressinus R. Br. ex G. Benn. var. Podocarpus guatemalensis Standl. var. allenii (Standl.)
curvulus Miq. 335 J. T. Buchholz & N. E. Gray 868
Podocarpus dacrydiifolius Wasscher 328 Podocarpus guatemalensis Standl. var. pinetorum
Podocarpus dacrydioides A. Rich. 327, 331 (Bartlett) J. T. Buchholz & N. E. Gray 868
Podocarpus dawei Stapf 141 Podocarpus hallii Kirk 852, 853
Podocarpus decipiens N. E. Gray 897 Podocarpus harmsianus Pilg. 948
Podocarpus decumbens N. E. Gray 825, 853 Podocarpus henkelii Stapf ex Dallim. & A.B. Jacks.
Podocarpus deflexus Ridl. 822, 854, 855 140, 812, 820, 870
Podocarpus degeneri (N. E. Gray) de Laub. ex Silba Podocarpus hispaniolensis de Laub. 827, 871
366, 897 Podocarpus humbertii de Laub. 820, 872
Podocarpus dispermus C. T. White 811, 825, 855, Podocarpus idenburgensis N. E. Gray 879
856, 984 Podocarpus imbricatus Blume 327, 334
Podocarpus distichus J. T. Buchholz 947 Podocarpus imbricatus Blume var. kinabaluensis
Podocarpus distichus J. T. Buchholz var. maialis Wasscher335
J. T. Buchholz 947 Podocarpus indonesiensis de Laub. & Silba 918, 919
Podocarpus ingensis de Laub. 901, 902 Podocarpus macrophyllus (Thunb.) Sweet var.
Podocarpus insularis de Laub. 873 piliramulus Z. X. Chen & Z. Q. Li 886
Podocarpus kawaii Hayata 334 Podocarpus macrostachyus Parl. 901, 902
Podocarpus koordersii Pilg. ex Koord. & Valeton Podocarpus madagascariensis Baker 820, 887
919 Podocarpus madagascariensis Baker var.
Podocarpus koshunensis (Kaneh.) Kaneh. 540 madagascariensis 888
Podocarpus ladei F. M. Bailey 949 Podocarpus madagascariensis Baker var. procerus
Podocarpus lambertii Klotzsch ex Endl. 196, 828 de Laub. 888
874, 875, 925, 938, 939 Podocarpus madagascariensis Baker var. rotundus
Podocarpus lambertii Klotzsch ex Endl. var. L. Laurent 888
1146 transiens Pilg. 938 Podocarpus magnifolius J. T. Buchholz & N. E.
Podocarpus latifolius Blume 143, 541, 812, 820, 829, Gray 827, 828, 845, 846, 889
876, Podocarpus maki (Siebold & Zucc.) Gaussen 886
Podocarpus latifolius (Thunb.) R. Br. ex Podocarpus mannii Hook. f. 145
Mirb. 876 Podocarpus matudae Lundell 813, 826, 829, 851,
Podocarpus latifolius (Thunb.) R. Br. ex Mirb. var. 890, 891, 1014
latior Pilg. 876 Podocarpus matudae Lundell var. jaliscanus de
Podocarpus latifolius Wall. 541 Laub. & Silba 890, 891
Podocarpus latior (Pilg.) Gaussen 876 Podocarpus matudae Lundell var. macrocarpus
Podocarpus laubenfelsii Tiong 360, 812, 821, 823, J.T. Buchholz & N.E. Gray 890, 891
877, 878 Podocarpus matudae Lundell var. reichei
Podocarpus lawrencei Hook. f. 374, 534, 561, 813, (J. T. Buchholz & N. E. Gray) de Laub. & Silba
825, 829, 867, 878, 879 890
Podocarpus ledermannii Pilg. 824, 879, 880 Podocarpus micropedunculatus de Laub. 823,
Podocarpus leonii Carabia 832 891, 892
Podocarpus leptophyllus Wasscher 362 Podocarpus milanjianus Rendle 146, 454, 813,
Podocarpus levis de Laub. 822, 823, 824, 880, 881 820, 871, 892, 893, 1061
Podocarpus longifoliolatus Pilg. 826, 881 Podocarpus minus (Carrire) Parl. 969
Podocarpus lophatus de Laub. 822, 882 Podocarpus montanus (Humb. & Bonpl. ex Willd.)
Podocarpus lucienii de Laub. 825, 883 Lodd. ex Britton 950
Podocarpus macrocarpus de Laub. 822, 883, 884 Podocarpus montanus (Willd.) Lodd. var. densifolius
Podocarpus macrophyllus (Thunb.) Sweet 175, J. T. Buchholz & N. E. Gray 950
273, 819, 821, 829, 846, 847, 862, 863, 884, 885, Podocarpus montanus (Willd.) Lodd. var.
893, 894, 1039, 1110 diversifolius Dallim. & A. B. Jacks. 950
Podocarpus macrophyllus (Thunb.) Sweet var. Podocarpus montanus (Willd.) Lodd. var. meridensis
chingii N. E. Gray 847, 885 J. T. Buchholz & N. E. Gray 950
Podocarpus macrophyllus (Thunb.) Sweet f. Podocarpus monteverdeensis de Laub. 901, 902
grandifolius Pilg. 862 Podocarpus nagi (Thunb.) Pilg. var. koshunensis
Podocarpus macrophyllus (Thunb.) Sweet var. Kaneh.540
liukiuensis Warb. 862, 863 Podocarpus nakaii Hayata 174, 489, 813, 821, 893,
Podocarpus macrophyllus (Thunb.) Sweet var. 894
macrophyllus 813, 886, 894 Podocarpus nankoensis Hayata 540
Podocarpus macrophyllus (Thunb.) Sweet subsp. Podocarpus neriifolius D. Don 175, 180, 182, 258,
maki (Siebold & Zucc.) Pilg. 886 277, 334, 821, 822, 823, 824, 826, 830, 836, 838,
Podocarpus macrophyllus (Thunb.) Sweet var. 868, 873, 874, 885, 895, 896, 897, 915, 920, 932,
maki Siebold & Zucc. 885, 886, 894 933, 973, 1017
Podocarpus macrophyllus (Thunb.) Sweet var. nakaii Podocarpus neriifolius D. Don var. atjehensis
(Hayata) H. L. Li & Keng 893 Wasscher836
Podocarpus neriifolius D. Don var. bracteatus Podocarpus pendulifolius J. T. Buchholz & N. E.
(Blume) Wasscher 838 Gray828, 905, 906, 907
Podocarpus neriifolius D. Don var. brevifolius Stapf Podocarpus perrieri Gaussen & Woltz 820, 907,
841 908
Podocarpus neriifolius D. Don var. decipiens Podocarpus philippinensis Foxw. 919
(N. E. Gray) Silba 897 Podocarpus pilgeri Foxw. 175, 180, 277, 821, 822,
Podocarpus neriifolius D. Don var. degeneri 823, 824, 832, 840, 908, 909, 914, 1008
N. E. Gray 895, 896, 897 Podocarpus pilgeri Foxw. var. thailandensis Gaussen
Podocarpus neriifolius D. Don var. 908
neriifolius 813, 896 Podocarpus pinetorum Bartlett 868, 869
Podocarpus neriifolius D. Don var. penibukanensis Podocarpus pittieri J. T. Buchholz & N. E. 1147
Silba897 Gray921
Podocarpus neriifolius D. Don var. polyanthus Podocarpus polyanthus (Wasscher) Gaussen 896
Wasscher896 Podocarpus polyspermus de Laub. 826, 910
Podocarpus neriifolius D. Don var. ridleyi Wasscher Podocarpus polystachyus R.Br. ex Endl. 814, 821,
915 822, 823, 824, 829, 850, 903, 910, 911, 912
Podocarpus neriifolius D. Don var. staintonii Silba Podocarpus polystachyus R. Br. ex Endl. var. rigidus
897 Wasscher837
Podocarpus neriifolius D. Don var. teysmannii Podocarpus polystachyus R. Br. ex Endl. var.
(Miq.) Wasscher 935 thevetiifolius (Zipp. ex Blume) Silba 910
Podocarpus neriifolius D. Don var. timorensis Podocarpus pseudobracteatus de Laub. 824, 912
Wasscher918 Podocarpus purdieanus Hook. 827, 913
Podocarpus nivalis Hook. 390, 391, 520, 814, 826, Podocarpus ramosii R. R. Mill 822, 823, 914, 915
829, 867, 897, 898 Podocarpus reichei J. T. Buchholz & N. E.
Podocarpus novae-caledoniae Vieill. 544, 814, Gray 890, 891
825, 898, 899, 933 Podocarpus ridleyi (Wasscher) N. E. Gray 822,
Podocarpus novae-caledoniae Vieill. var. colliculatus 915, 927
N. E. Gray 933 Podocarpus roraimae Pilg. 828, 916, 917, 930, 935
Podocarpus nubigenus Lindl. 382, 625, 814, 828, Podocarpus rospigliosii Pilg. 971
829, 900, 901, 976 Podocarpus rostratus L. Laurent 820, 845, 908,
Podocarpus oleifolius D. Don 827, 828, 846, 869, 917
901, 902 Podocarpus rostratus L. Laurent var. perrieri
Podocarpus oleifolius D. Don var. costaricensis (Gaussen & Woltz) Silba 907
J. T. Buchholz & N. E. Gray 901 Podocarpus rotundus de Laub. 914
Podocarpus oleifolius D. Don var. equadorensis Podocarpus rubens de Laub. 918, 919
Silba901 Podocarpus rubens de Laub. var. pabinamaensis
Podocarpus oleifolius D. Don var. macrostachyus Silba918
(Parl.) J. T. Buchholz & N. E. Gray 901 Podocarpus rubens de Laub. var. sumatrana Silba
Podocarpus oleifolius D. Don var. trujillensis 918
J. T. Buchholz & N. E. Gray 901 Podocarpus rumphii Blume 814, 821, 823, 824,
Podocarpus orarius R. R. Mill & M. Whiting 1073 884, 919, 920
Podocarpus palawanensis de Laub. & Silba 822, Podocarpus rumphii Blume var. arbainii Silba 919
902 Podocarpus rumphii Blume var. aruensis Silba 919
Podocarpus pallidus N. E. Gray 826, 903, 904 Podocarpus rusbyi J. T. Buchholz & N. E.
Podocarpus palustris J. T. Buchholz 969 Gray827, 920, 921
Podocarpus papuanus Ridl. 329, 335 Podocarpus salicifolius Klotzsch & H. Karst. ex
Podocarpus parlatorei Pilg. 828, 904, 905, 921 Endl.827, 906, 921, 922
Podocarpus pectinatus Pancher ex Brongn. & Gris Podocarpus salignus D. Don 382, 815, 828, 829,
133 922, 923
Podocarpus salomoniensis Wasscher 824, 923 Podocarpus utilior Pilg. 948, 949
Podocarpus sellowii Klotzsch ex Endl. 196, 827, Podocarpus victorinianus Carabia 832, 833
874, 924, 925 Podocarpus vieillardii Parl. 338
Podocarpus sellowii Klotzsch ex Endl. var. Podocarpus vitiensis Seem. 967, 972
angustifolius Pilg. 926 Podocarpus wallichianus C. Presl 541
Podocarpus sellowii Klotzsch ex Endl. var. sellowii Podocarpus woltzii Gaussen 844, 845
925 Prumnopitys Phil. 20, 43, 53, 54, 943, 949, 953,
Podocarpus smithii de Laub. 825, 926, 927, 950 972, 984
Podocarpus spathoides de Laub. 822, 823, 824, Prumnopitys amara (Blume) de Laub. 983
927, 928 Prumnopitys andina (Poepp. ex Endl.) de Laub.
1148 Podocarpus spathoides de Laub. var. solomonensis 382, 816, 943, 944, 945
Silba1071 Prumnopitys andina (Poepp. ex Endl.) de Laub.
Podocarpus spicatus Poepp. 944 subsp. blijdensteinii Silba 944
Podocarpus spinulosus (Sm.) R. Br. ex Mirb. 815, Prumnopitys elegans Phil. 943, 944
825, 928 Prumnopitys exigua de Laub. ex Silba 943, 945, 953
Podocarpus sprucei Parl. 815, 828, 929 Prumnopitys ferruginea (G. Benn. ex D. Don) de
Podocarpus standleyi J. T. Buchholz & N. E. Gray Laub. 152, 332, 356, 390, 392, 525, 566, 853, 937,
951 944, 946, 947
Podocarpus steupii Wasscher 337 Prumnopitys ferruginoides (R. H. Compton) de
Podocarpus steyermarkii J. T. Buchholz & N. E. Laub. 208, 211, 363, 522, 816, 947, 948
Gray 827, 917, 930, 931, 935 Prumnopitys harmsiana (Pilg.) de Laub. 944,
Podocarpus subtropicalis de Laub. 821, 830, 895, 948, 949
896, 932 Prumnopitys ladei (F.M. Bailey) de Laub. 816,
Podocarpus subtropicalis de Laub. var. medogensis 949, 950
Silba932 Prumnopitys montana (Humb. & Bonpl. ex Willd.)
Podocarpus sylvestris J. T. Buchholz 161, 221, de Laub. 944, 946, 950, 951
933 Prumnopitys spicata (Poepp.) Molloy & Muoz-
Podocarpus tepuiensis J. T. Buchholz & N. E. Gray Schick944
828, 917, 930, 934, 935 Prumnopitys standleyi (J. T. Buchholz & N. E.
Podocarpus teysmannii Miq. 821, 823, 935, 936 Gray) de Laub. 951, 952
Podocarpus thevetiifolius Zipp. ex Blume 910 Prumnopitys taxifolia (Banks & Sol. ex D. Don) de
Podocarpus tixieri Gaussen 908 Laub. 152, 332, 356, 390, 394, 525, 566, 943, 947,
Podocarpus tixieri Gaussen ex Silba 908 952, 953
Podocarpus totara G. Benn. ex D. Don 20, 28, Pseudolarix Gordon 17, 19, 41, 51, 52, 954
143, 152, 332, 356, 525, 566, 815, 826, 829, 853, Pseudolarix amabilis (J. Nelson) Rehd. 280, 816,
936, 937, 947, 953 954, 955
Podocarpus totara G. Benn. ex D. Don var. hallii Pseudolarix kaempferi (Lindl.) Gordon 954
(Kirk) Pilg. 852 Pseudolarix pourtetii Ferr 954
Podocarpus totara G. Benn. ex D. Don var. Pseudotaxus W. C. Cheng 44, 55, 957
waihoensis Wardle 936 Pseudotaxus chienii (W. C. Cheng) W. C. Cheng
Podocarpus transiens (Pilg.) de Laub. ex 297, 816, 957, 958
Silba828, 938 Pseudotaxus chienii (W.C. Cheng) W. C. Cheng
Podocarpus transiens (Pilg.) de Laub. ex Silba var. subsp. liana (Silba) Silba 957
harleyi Silba 938 Pseudotaxus liana Silba 957
Podocarpus trinitensis J. T. Buchholz & N. E. Gray Pseudotsuga Carrire 19, 40, 52, 259, 652, 678,
827, 939, 940 708, 785, 959, 965
Podocarpus urbanii Pilg. 826, 940, 941, 942 Pseudotsuga argyrophylla (Chun & Kuang) Greguss
Podocarpus usambarensis Pilg. 146 259
Podocarpus ustus (Vieill.) Brongn. & Gris 555 Pseudotsuga brevifolia W. C. Cheng & L. K. Fu 965
Pseudotsuga davidiana Bertrand 490 Retinispora lawsoniana (A. Murray bis) A. V.
Pseudotsuga flahaultii Flous 963 Bobrov & Melikyan 283
Pseudotsuga forrestii Craib 965 Retinispora obtusa Siebold & Zucc. 281, 286
Pseudotsuga gausseni Flous 966 Retinispora pisifera Siebold & Zucc. 288
Pseudotsuga japonica (Shiras.) Beissn. 86, 286, Retinispora taiwanensis (Masam. & S. Suzuki)
541, 960, 993, 1055 A. V. Bobrov & Melikyan 286
Pseudotsuga macrocarpa (Vasey) Mayr 254, 959, Retrophyllum C. N. Page 42, 53, 54, 967
961 Retrophyllum comptonii (J. T. Buchholz)
Pseudotsuga menziesii (Mirb.) Franco 27, 52, 63, C. N. Page 166, 967, 968, 994
70, 91, 102, 117, 254, 284, 300, 304, 313, 322, 510, Retrophyllum minus (Carrire) C. N. Page 20,
581, 584, 617, 690, 697, 701, 719, 727, 737, 772, 350, 353, 361, 967, 969, 970, 994 1149
959, 962, 980, 982, 993, 1005, 1026, 1033, 1052 Retrophyllum piresii (Silba) C. N. Page 967, 970
Pseudotsuga menziesii (Mirb.) Franco var. flahaultii Retrophyllum rospigliosii (Pilg.) C. N. Page 967,
(Flous) Silba 963 971, 972
Pseudotsuga menziesii (Mirb.) Franco subsp. glauca Retrophyllum vitiense (Seem.) C. N. Page 835,
(Beissn.) E. Murray 963 967, 972, 973, 974
Pseudotsuga menziesii (Mirb.) Franco var. glauca
(Beissn.) Franco 76, 80, 81, 120, 129, 317, 582, S
611, 963, 1026
Pseudotsuga menziesii (Mirb.) Franco subsp. Sabina Mill.393
macrocarpa (Vasey) E. Murray 961 Sabina ashei (J. T. Buchholz) A. V. Bobrov &
Pseudotsuga menziesii (Mirb.) Franco var. Melikyan400
menziesii 963, 993 Sabina convallium (Rehd. & E. H. Wilson)
Pseudotsuga menziesii (Mirb.) Franco var. oaxacana W. C. Cheng & L. K. Fu var. microsperma
Debreczy & Rcz 963 W. C. Cheng & L. K. Fu 418
Pseudotsuga shaanxiensis S. Z. Qu & K. Y. Wang Sabina microsperma (W. C. Cheng & L. K. Fu)
965 W. C. Cheng & L. K. Fu 418
Pseudotsuga sinensis Dode 98, 182, 255, 258, 277, Sabina pingii (W. C. Cheng) W. C. Cheng &
295, 492, 537, 541, 604, 885, 964, 986, 1008, W. T. Wang var. wilsonii (Rehd.) W. C. Cheng
1050, 1068 & L. K. Fu 452, 453
Pseudotsuga sinensis Dode var. brevifolia (W. C. Sabina silicicola Small486
Cheng & L. K. Fu) Farjon & Silba 965 Sabina vulgaris Antoine393
Pseudotsuga sinensis Dode var. forrestii (Craib) Sabina vulgaris Antoine var. erectopatens
Silba965 W. C. Cheng & L. K. Fu 409
Pseudotsuga sinensis Dode var. gaussenii (Flous) Sabina vulgaris Antoine var. yulinensis T. C. Chang
Silba959, 966 & C. G. Chen 463
Pseudotsuga sinensis Dode var. sinensis 959, 965, Sabinella Nakai393
993 Sabinella phoenicea (L.) Nakai 393
Pseudotsuga sinensis Dode var. wilsoniana (Hayata) Salisburyodendron A. V. Bobrov & Melikyan 148
L. K. Fu & Nan Li 965 Salisburyodendron australis (D. Don) A. V. Bobrov
Pseudotsuga wilsoniana Hayata 965 & Melikyan 148, 151
Pseudotsuga xichangensis C. T. Kuan & L. J. Zhou Salisburyodendron corbassonii (de Laub.) A. V.
965 Bobrov & Melikyan 166
Salisburyodendron lanceolata (Warb.) A. V. Bobrov
R & Melikyan 161
Salisburyodendron montana (de Laub.) A. V. Bobrov
Retinispora Siebold & Zucc. 281 & Melikyan 166
Retinispora formosensis (Matsum.) A. V. Bobrov & Salisburyodendron moorei (Lindl.) A. V. Bobrov &
Melikyan281 Melikyan166
Salisburyodendron ovata (Vieill.) A. V. Bobrov & Taiwania flousiana Gaussen985
Melikyan169 Taiwania yunnanensis Koidz.985
Salisburyodendron ovata (Vieill.) A. V. Bobrov Tassilicyparis A. V. Bobrov & Melikyan 298
& Melikyan subsp. hypoleuca (C. Moore Tassilicyparis dupreziana (A. Camus) A. V. Bobrov
ex Henkel & W. Hochst.) A. V. Bobrov & & Melikyan 298, 310
Melikyan169 Taxaceae Gray 18, 21, 22, 24, 26, 35, 43, 45, 47, 55,
Saxegothaea Lindl. 41, 53, 54, 975 174, 228, 229, 258, 555, 957, 1001, 1032, 1094,
Saxegothaea conspicua Lindl. 382, 900, 975, 976, 1096, 1099, 1100, 1102
994 Taxodium Rich. 17, 21, 37, 48, 49, 388, 953, 988
Schubertia Mirb.988 Taxodium ascendens Brongn.990
1150 Schubertia disticha (L.) Mirb. 988 Taxodium distichum (L.) Rich. 290, 322, 531, 691,
Sciadopityaceae Luerss. 21, 24, 43, 45, 54, 977, 978, 700, 782, 979, 988, 989, 990, 991, 996
1074 Taxodium distichum (L.) Rich. var. ascendens
Sciadopitys Siebold & Zucc. 18, 43, 48, 54, 317, (Brongn.) Mast. 990
393, 436, 452, 464, 624, 977, 978 Taxodium distichum (L.) Rich. var. distichum 990
Sciadopitys verticillata (Thunb.) Siebold & Zucc. Taxodium distichum (L.) Rich. var. imbricarium
21, 86, 286, 977, 994, 1030, 1039, 1055 (Nutt.) Sarg. 990
Sequoia Endl. 21, 24, 36, 48, 49, 238, 530, 979 Taxodium distichum (L.) Rich. var. mucronatum
Sequoia sempervirens (D. Don) Endl. 21, 48, 284, (Ten.) A. Henry 990
979, 980, 995, 1005, 1026, 1033, 1077 Taxodium distichum (L.) Rich. subsp. nutans
Sequoiadendron J. T. Buchholz 24, 36, 48, 49, 238, (Aiton) E. Murray 990
282, 981 Taxodium mucronatum Ten.988, 990, 991, 996
Sequoiadendron giganteum (Lindl.) J. T. Buchholz Taxodium sempervirens D. Don 979
21, 25, 254, 546, 711, 719, 981, 982, 995 Taxus L. 22, 28, 44, 55, 174, 175, 268, 380, 481, 511,
Squamataxus J. Nelson 975 555, 562, 975, 978, 1001, 1009, 1013, 1014, 1016,
Squamataxus albertiana J. Nelson 975 1017, 1040
Stachycarpus Tiegh.943 Taxus baccata L. 74, 111, 113, 263, 610, 620, 757,
Stachycarpus amarus (Blume) Gaussen 983 996, 997, 1001, 1002, 1003, 1004, 1005, 1007,
Stachycarpus andinus (Poepp. ex Endl.) Tiegh. 1011, 1017, 1018
943 Taxus baccata L. subsp. brevifolia (Nutt.)
Stegocedrus Doweld521 Pilg.1004
Stegocedrus austrocaledonica (Brongn. & Gris) Taxus baccata L. subsp. canadensis (Marshall) Pilg.
Doweld 521, 522 1005
Stegocedrus chevalieri (J. T. Buchholz) Doweld 524 Taxus baccata L. subsp. cuspidata (Siebold & Zucc.)
Stegocedrus yateensis (Guillaumin) Doweld 526 Pilg.1011
Steinhauera Presl981 Taxus baccata L. subsp. cuspidata (Siebold & Zucc.)
Steinhauera gigantea (Lindl.) Kuntze ex Voss981 Pilg. var. chinensis Pilg.1007
Strobus Opiz626 Taxus baccata L. subsp. cuspidata (Siebold & Zucc.)
Strobus weymouthiana Opiz626 Pilg. var. latifolia Pilg.1011
Sundacarpus (J. T. Buchholz & N. E. Gray) Taxus baccata L. subsp. floridana (Nutt. ex Chapm.)
C. N. Page 43, 53, 54, 378, 983 Pilg.1012
Sundacarpus amarus (Blume) C. N. Page 155, 331, Taxus baccata L. var. microcarpa Trautv.1011
352, 360, 377, 378, 542, 878, 950, 983, 984, 995 Taxus baccata L. var. minor Michx.1005
Taxus baccata L. subsp. wallichiana (Zucc.) Pilg.
T 1016, 1017
Taxus biternata Spjut1011
Taiwania Hayata 36, 48, 49, 985, 986 Taxus brevifolia Nutt. 284, 737, 982, 997, 1002,
Taiwania cryptomerioides Hayata 255, 282, 286, 1004, 1005, 1026
295, 985, 986, 995, 996 Taxus brevifolia Nutt. var. polychaeta Spjut1004
Taxus brevifolia Nutt. var. reptaneta Spjut1004, Taxus kingstonii Spjut1014
1005 Taxus latifolia Thunb.876
Taxus caespitosa Nakai1011 Taxus macrophylla Thunb. 819, 886
Taxus caespitosa Nakai var. angustifolia Spjut1011 Taxus mairei (Leme & Lv.) S. Y. Hu ex T. S. Liu
Taxus caespitosa Nakai var. latifolia (Pilg.) Spjut 1002, 1014, 1015, 1017
1011 Taxus mairei (Leme & Lv.) S. Y. Hu ex T. S. Liu
Taxus canadensis Marshall 65, 1002, 1004, 1005, var. speciosa (Florin) Spjut 1014
1006 Taxus media Rehd.1011
Taxus canadensis Marshall var. floridana (Nutt. ex Taxus minor (Michx.) Britton ex Small & Vail
Chapm.) Silba 1012 1005
Taxus canadensis Marshall var. minor (Michx.) Taxus montana Humb. & Bonpl. ex Willd.950 1151
Spjut1006 Taxus nucifera L. 1032, 1038
Taxus celebica (Warb.) H. L. Li 1016, 1017 Taxus obscura Spjut1016
Taxus chienii W. C. Cheng 957 Taxus phytonii Spjut1016
Taxus chinensis (Pilg.) Rehd. 66, 84, 175, 182, 270, Taxus speciosa Florin 1014, 1015
277, 297, 492, 541, 885, 1007, 1014, 1016, 1017, Taxus spinulosa Sm.928
1068 Taxus suffnessii Spjut1016
Taxus chinensis (Pilg.) Rehd. var. mairei (Leme & Taxus sumatrana (Miq.) de Laub. 1016, 1017
Lv.) W. C. Cheng & L. K. Fu 1014 Taxus umbraculifera (Siebold ex Endl.) Lawson var.
Taxus chinensis (Pilg.) Rehd. var. yunnanensis microcarpa (Trautv.) Spjut 1011
(W. C. Cheng & L. K. Fu) L. K. Fu 1016 Taxus umbraculifera (Siebold ex Endl.) Lawson var.
Taxus communis J. Nelson 1001 nana (Rehd.) Spjut 1011
Taxus contorta Griff.1002, 1008, 1009, 1017, 1018 Taxus verticillata Thunb.977
Taxus contorta Griff. var. mucronata Spjut1016 Taxus wallichiana Zucc. 277, 579, 620, 986, 1002,
Taxus cuspidata Siebold & Zucc. 100, 590, 1002, 1007, 1009, 1015, 1016, 1017, 1018
1007, 1009, 1010, 1011, 1039 Taxus wallichiana Zucc. var. chinensis (Pilg.)
Taxus cuspidata Siebold & Zucc. var. caespitosa Florin1007
(Nakai) Q. L. Wang 1011 Taxus wallichiana Zucc. var. mairei (Leme & Lv.)
Taxus cuspidata Siebold & Zucc. var. chinensis L. K. Fu & Nan Li 1014
(Pilg.) C. K. Schneid. 1007 Taxus wallichiana Zucc. var. yunnanensis
Taxus cuspidata Siebold & Zucc. var. (W. C. Cheng & L. K. Fu) C. T. Kuan 1016
cuspidata997, 1011 Taxus yunnanensis W. C. Cheng & L. K. Fu 1009,
Taxus cuspidata Siebold & Zucc. var. latifolia (Pilg.) 1016, 1036, 1050
Nakai1011 Tetraclinis Mast. 38, 48, 49, 532, 1019, 1021
Taxus cuspidata Siebold & Zucc. var. microcarpa Tetraclinis articulata (Vahl) Mast. 28, 289, 310,
(Trautv.) Kolesn. 1011 997, 1019, 1020
Taxus cuspidata Siebold & Zucc. var. nana hort. ex Thalamia Spreng.560
Rehd. 1011 Thalamia aspleniifolia (Labill.) Spreng. 560
Taxus elongata Aiton 819, 860 Thuja L. 37, 48, 50, 226, 532, 817, 963, 1022, 1030
Taxus falcata Thunb. 140, 142 Thuja articulata Vahl1019
Taxus floridana Nutt. ex Chapm.1002, 1012, 1013, Thuja chengii 817
1040, 1041 Thuja chilensis D. Don 226
Taxus florinii Spjut1016 Thuja cupressoides L. 1056, 1058
Taxus fuana Nan Li & R. R. Mill 1008, 1009, 1017 Thuja dolabrata Thunb. ex L. f. 1030, 1031
Taxus globosa Schltdl. 601, 1002, 1012, 1013, 1014 Thuja koraiensis Nakai 997, 998, 1022, 1023
Taxus globosa Schltdl. var. floridana (Nutt. ex Thuja lineata Poir.387
Chapm.) Spjut 1012 Thuja occidentalis L. 804, 1022, 1024, 1026
Taxus harringtonii Knight ex J. Forbes 268, 272, Thuja orientalis L.817
273 Thuja pensilis Staunton ex D. Don 387
Thuja plicata Donn ex D. Don 63, 91, 117, 254, Torreya yunnanensis W. C. Cheng & L. K. Fu
510, 617, 719, 737, 998, 1005, 1022, 1025, 1026, 1035
1066 Tsuga (Endl.) Carrire 19, 41, 52, 273, 274, 432,
Thuja sphaeroidea Spreng.281 459, 549, 551, 573, 595, 954, 959, 963, 1031, 1042
Thuja standishii (Gordon) Carrire 97, 128, 1022, Tsuga argyrophylla (Chun & Kuang) de Laub. &
1027, 1030 Silba259
Thuja sutchuenensis Franch. 998, 1022, 1028 Tsuga blaringhemii Flous1047
Thujopsis Siebold & Zucc. ex Endl. 37, 48, 49, 1030 Tsuga canadensis (L.) Carriere 64, 615, 775, 804,
Thujopsis dolabrata (Thunb. ex L. f.) Siebold & 1006, 1024, 1042, 1043, 1044, 1046
Zucc.1027, 1030 Tsuga caroliniana Engelm. 90, 1042, 1043, 1044,
1152 Thujopsis dolabrata (Thunb. ex L. f.) Siebold & 1045, 1046
Zucc. var. dolabrata998, 1031 Tsuga chinensis (Franch.) E. Pritz. 66, 72, 77, 82,
Thujopsis dolabrata (Thunb. ex L. f.) Siebold & 83, 84, 118, 131, 180, 182, 258, 260, 282, 511, 551,
Zucc. var. hondae Makino 1031 578, 604, 605, 612, 614, 885, 965, 986, 1008,
Thujopsis standishii Gordon1027 1042, 1045, 1046, 1050
Titanodendron A. V. Bobrov & Melikyan 191 Tsuga chinensis (Franch.) E. Pritz. var. chinensis
Titanodendron hunsteinii (K. Schum.) A. V. Bobrov 98, 999, 1046
& Melikyan 191, 208 Tsuga chinensis (Franch.) E. Pritz. var. daibuensis
Titanodendron klinkii (Lauterb.) A. V. Bobrov & S. S. Ying 1047
Melikyan208 Tsuga chinensis (Franch.) E. Pritz. var. formosana
Titanodendron schumanniana (Warb.) A. V. Bobrov (Hayata) H. L. Li & H. Keng 1046
& Melikyan 208 Tsuga chinensis (Franch.) E. Pritz. var. forrestii
Torreya Arn. 22, 44, 55, 1032, 1040 (Downie) Silba 1050
Torreya ascendens Nakai ex Uyeki1038 Tsuga chinensis (Franch.) E. Pritz. var.
Torreya californica Torr. 998, 1032, 1033 oblongisquamata W. C. Cheng & L. K. Fu
Torreya fargesii Franch.1032, 1034 1047
Torreya fargesii Franch. var. fargesii 1034, 1035 Tsuga chinensis (Franch.) E. Pritz. subsp. patens
Torreya fargesii Franch. var. yunnanensis (Downie) E. Murray 1046
(W. C. Cheng & L. K. Fu) N. Kang 1034, 1035 Tsuga chinensis (Franch.) E. Pritz. var. patens
Torreya grandis Fortune ex Lindl. 297, 489, 1032, (Downie) L. K. Fu & Nan Li 1046
1034, 1035, 1036 Tsuga chinensis (Franch.) E. Pritz. var. robusta
Torreya grandis Fortune ex Lindl. var. fargesii W. C. Cheng & L. K. Fu 1047
(Franch.) Silba 1035 Tsuga chinensis (Franch.) E. Pritz. var.
Torreya grandis Fortune ex Lindl. var. grandis tchekiangensis (Flous) W. C. Cheng &
1036, 1037 L. K. Fu 1046
Torreya grandis Fortune ex Lindl. var. Tsuga chinensis (Franch.) E. Pritz. subsp. wardii
jiulongshanensis Z. Y. Li, Z. C. Tang & (Downie) E. Murray 1049
N. Kang 1034, 1037 Tsuga diversifolia (Maxim.) Mast. 97, 105, 128,
Torreya grandis Fortune ex Lindl. var. yunnanensis 506, 573, 591, 1027, 1030, 1042, 1043, 1047, 1048
(W. C. Cheng & L. K. Fu) Silba 986 Tsuga diversifolia (Maxim.) Mast. subsp.
Torreya jackii Chun1032, 1037, 1038 blaringhemii (Flous) E. Murray 1047
Torreya nucifera (L.) Siebold & Zucc. 86, 293, Tsuga douglasii (Sabine ex D. Don) Lindl. var.
960, 999, 1032, 1038, 1039 glauca Beissn.963
Torreya nucifera (L.) Siebold & Zucc. var. grandis Tsuga dumosa (D. Don) Eichler 77, 78, 79, 87, 114,
(Fortune ex Lindl.) Pilg. 1037 125, 305, 504, 511, 579, 586, 618, 620, 805, 965,
Torreya parvifolia T. P. Yi, L. Yang & T. L. Long 986, 1043, 1048, 1049, 1050
1037 Tsuga dumosa (D. Don) Eichler subsp. leptophylla
Torreya taxifolia Arn. 1013, 1032, 1040, 1093, 1101 (Hand.-Mazz.) E. Murray 1049
Tsuga dumosa (D. Don) Eichler var. yunnanensis W
(Franch.) Silba 1048
Tsuga dura Downie1048 Washingtonia Winslow 981, 982
Tsuga formosana Hayata1046 Washingtonia californica Winslow 981
Tsuga forrestii Downie126, 999, 1042, 1043, 1050, Wellingtonia Lindl. 981, 982
1051 Wellingtonia gigantea Lindl. 981, 982
Tsuga heterophylla (Raf.) Sarg. 63, 91, 117, 254, Widdringtonia Endl. 24, 39, 48, 50, 1056
284, 510, 598, 617, 719, 737, 980, 999, 1005, 1026, Widdringtonia cedarbergensis J. A. Marsh 999,
1042, 1043, 1051, 1052, 1053, 1054, 1066 1056, 1100
Tsuga japonica Shiras.960 Widdringtonia cupressoides (L.) Endl. 1058
Tsuga jeffreyi (A. Henry) A. Henry 1042, 1054 Widdringtonia dracomontana Stapf 1058 1153
Tsuga leptophylla Hand.-Mazz.1049 Widdringtonia nodiflora (L.) Powrie 1000, 1056,
Tsuga longibracteata W. C. Cheng 549, 1042 1058, 1061
Tsuga mairei Leme & Lv. 1014 Widdringtonia nodiflora (L.) Powrie var.
Tsuga mertensiana (Bong.) Carrire 63, 101, 117, dracomontana (Stapf) Silba 1058
443, 508, 510, 581, 598, 617, 637, 674, 737, 1026, Widdringtonia nodiflora (L.) Powrie var. whytei
1042, 1052, 1053, 1054, 1066 (Rendle) Silba 1060
Tsuga mertensiana (Bong.) Carrire subsp. Widdringtonia schwarzii (Marloth) Mast. 1056,
grandicona Farjon 102, 1043, 1054 1059, 1060
Tsuga mertensiana (Bong.) Carrire subsp. Widdringtonia whytei Rendle 1000, 1056, 1060,
mertensiana 1043, 1053 1061
Tsuga mertensiana (Bong.) Carrire subsp. Wollemia W. G. Jones et al. 18, 21, 35, 46, 1062,
mertensiana var. jeffreyi (A. Henry) 1063
C. K. Schneid. 1042, 1054 Wollemia nobilis W. G. Jones et al.18, 1000, 1062,
Tsuga mertensiana (Bong.) Carrire subsp. 1097
mertensiana var. mertensiana999, 1053
Tsuga oblongisquamata (W. C. Cheng & L. K. Fu) X
L. K. Fu & Nan Li 1047
Tsuga patens Downie1046 Xanthocyparis Farjon & Hiep 38, 48, 49, 298, 299,
Tsuga pattoniana (Balf.) Senecl. var. jeffreyi 1064, 1065, 1068
A. Henry 1042, 1054 Xanthocyparis nootkatensis (D. Don) Farjon &
Tsuga sieboldii Carrire 86, 286, 293, 541, 960, Harder617, 1000, 1026, 1053, 1065, 1066
978, 1039, 1042, 1043, 1054, 1055 Xanthocyparis vietnamensis Farjon & Hiep 180,
Tsuga tchekiangensis Flous1046 182, 258, 1000, 1008, 1064, 1065, 1066, 1067,
Tsuga wardii Downie1048 1093, 1096
Tsuga yunnanensis (Franch.) E. Pritz. 1048
Tsuga yunnanensis (Franch.) E. Pritz. subsp. dura End notes
(Downie) E. Murray 1048
Tumion Raf. ex Greene1032 *The following new combination has been pub-
Tumion taxifolium (Arn.) Greene 1032 lished in this book: