An Atlas of the World’s Conifers

An Atlas of the World’s Conifers

An Analysis of their Distribution, Biogeography, Diversity

and Conservation Status

By

Aljos Farjon and Denis Filer

Leiden • boston
2013
front cover:
Top: Map of the world with distribution of all conifers; the shaded boreal zone represents broadly
contiguous conifer coverage
Middle: Larix sibirica and Picea abies in Siberian taiga (photo Bryan Alexander)
Bottom left to right: Pinus hwangshanensis, Sequoia sempervirens, Araucaria araucana
(photos Aljos Farjon)

back cover:
Maps left to right: Afrocarpus gracilior, conifers in South America, genus Phyllocladus, genus Pinus
in Mexico (partly), Podocarpus totara
Photos clockwise from top: Pinus hartwegii, Picea schrenkiana, Pinus balfouriana
(photos Aljos Farjon)

Library of Congress Cataloging-in-Publication Data

Farjon, Aljos.
 An atlas of the world’s conifers : an analysis of their distribution, biogeography, diversity, and
conservation status / by Aljos Farjon and Denis Filer.
 pages cm
 Includes bibliographical references.
 ISBN 978-90-04-21180-3 (hardback : alk. paper) — ISBN 978-90-04-21181-0 (e-book)
1. Conifers—Geographical distribution. I. Filer, Denis. II. Title.

 QK494.F354 2013
 585—dc23
2013022235

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ISBN 978 90 04 21180 3 (hardback)

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 Contents

Foreword ......................................................................................................................................................... vii
Preface .............................................................................................................................................................. ix
Acknowledgements ...................................................................................................................................... xi

Introduction ................................................................................................................................................... 1

Creating the Atlas of the World’s Conifers ........................................................................................... 3

Global and Trans-Continental Distributions ........................................................................................ 11

North America ............................................................................................................................................... 34

Mexico, Central America and the West Indies .................................................................................... 90

Europe and the Mediterranean ................................................................................................................ 139

Mainland Asia and Japan ........................................................................................................................... 171

Malesia ............................................................................................................................................................. 293

Australasia ....................................................................................................................................................... 339

South America ............................................................................................................................................... 428

Africa ................................................................................................................................................................ 458

Examples of Further Research .................................................................................................................. 474

References ....................................................................................................................................................... 489

Tables and Indices ........................................................................................................................................ 494

 Foreword

Aljos Farjon has done so much to provide comprehensive information about the conifers and
Denis Filer has provided a data management system for botanists, which is now so widely used
around the world. What an excellent combination to bring these two specialists together to pro-
duce An Atlas of the World’s Conifers. This first comprehensive Atlas for the conifers is a logical
sequence to Farjon’s landmark publication A Handbook of the World’s Conifers. Also available is
the conifer website providing interactive access to all the data used to create the Atlas maps, an
invaluable resource for those working on conifer biogeography.
Atlases are always interesting and this one is no exception. It is based on such a thorough
study that is backed up by a database of some 37,000 records. It is wonderful to have a work
that presents maps of every living species of conifer (615 in all) together with insightful analyses
of the patterns of distribution. This is much more than a collection of species maps. The analy-
ses accompanying the maps contain much useful information about the geography, climate and
vegetation of each major region of the world and have much useful biogeographic information
and comparisons between regions. In addition to the maps for individual species are the regional
maps and those that analyse the numbers of species, genera and families by degree cell.
This is yet another publication that demonstrates the importance of the herbaria of the world.
Herbaria are a treasure of information about plants and it is good to see this being put to good
use here. The information to produce this Atlas is based on data from a large number of geo-
referenced herbarium specimens from many of the world’s herbaria. This is the sort of work that
is an invaluable tool for conservation. Many conifers are quite rare and have restricted distribu-
tions and I hope that this book will not just remain on library shelves but will be put to use for
conservation planning. The fact that the contents of this work are available electronically as well
as in print means that there are no excuses for delaying conservation action. I compliment the
authors for this most valuable and interesting new resource on conifers.

Ghillean T. Prance FRS

former Director of the Royal Botanic Gardens, Kew
 Preface

The main content of this Atlas of the World’s Conifers is the distribution maps. There are 728 maps
in the Atlas. There are separate maps for all families, for all genera with more than one species,
for nearly all species (only a few are combined), while subspecies and varieties, when shown, are
on the map of the relevant species with different symbols. In addition to these 656 taxonomic
maps, there are 72 additional maps, ranging from a global map of all conifer species to continental
maps with the same, maps of families or genera or all species in parts of their range, e.g. China,
Japan, Mexico, USA, maps that analyse taxonomic diversity distribution at genus and species +
infraspecific levels, maps that calculate occurrence of rarity and other thematic maps. The main
part of the Atlas is formed by the chapters that deal with the taxonomic distribution of conifers,
beginning with Global and trans-continental distributions and proceeding through first the North-
ern Hemisphere with North America; Mexico, Central America and the West Indies; Europe and the
Mediterranean; Mainland Asia and Japan; then through the Tropics and the Southern Hemisphere
with Malesia; Australasia; South America and Africa. Genera and species have been given a map in
the chapter where they appear to fit best; e.g. the species that occur in Algeria and Morocco N of
the Sahara are not treated in the chapter Africa but in the chapter Europe and the Mediterranean.
Truly African conifers occur almost all S of the Sahara and most of the species N of the Sahara
either occur also in Europe or are very closely related to species that do. Similarly, species that
cross over into the area of another chapter have been given a map where most of the distribu-
tion is situated. This is usually explained in the text at a point where the reader might expect a
map to be given but is not. The indices at the end of the Atlas give the map numbers and pages
where each map can be found. Maps are numbered by chapter and the chapters are indicated
by an acronym.
The distribution of a taxon or taxa is represented on a map in map points; each point indi-
cates the approximate locality where one or more herbarium collections of that taxon or those
taxa have been made. For a discussion of the methods of data gathering and mapping and the
nature and quality of the data we refer the reader to the chapter Creating the Atlas of the World’s
Conifers. Here we need to point out what the maps do and do not show, or in other words, how
to read them. The maps show localities of herbarium collections made in the past. With each
species map there are figures given in brackets that give the following data: ‘n’ is the number of
herbarium collections (records) in the Conifer Database; ‘m’ is the number of ‘mappable’ records,
i.e. those for which locality coordinates in degrees and minutes could be given; ‘h’ is the number
of institutional herbaria in which the specimens (unicates and duplicates of collections) are kept;
this is followed by the period in which these collections were made. These statistics are indica-
tors of how complete or incomplete the data used for the maps are. The map points are together
a subset of the presence of a species, limited to those trees from which a sample was taken for
a herbarium. No other data sources were used. The maps should be read with these limitations
in mind.
The text accompanies the maps and gives factual information on distribution, habitat, abun-
dance or rarity, the conservation status if a species falls within a threatened IUCN Red List category
and sometimes other topics. There may also be explanatory text on for example completeness
of records, comparisons with other taxa and interpretation of the distribution patterns. The tax-
onomy refers to A Handbook of the World’s Conifers by the first author, published by Brill in 2010,
and remains unaltered in this Atlas. In addition to the text with the maps, each chapter has one
or more introductory sections that outline the general geography of a continent, or smaller area,
x preface

if it is of particular note to conifer distribution (e.g. China, Japan, New Zealand). This information
is kept as brief as possible and should not be read as an essay on the geography of that continent
or country. These introductions serve merely to put in geographical perspective that which fol-
lows, which is an outline of conifer distribution and diversity in that area. Maps are here again
supreme, and the text accompanies these and describes and interprets the distribution patterns.
The first map in the chapter or section gives the total distribution of all species in the area. These
and other maps like them are based on all records of species and their subspecies and varieties
that occur there. They show not only where conifer species occur, but also where they are con-
centrated. The latter aspect is further analysed in diversity maps, usually at genus and species +
infraspecific levels in two separate maps, calculated for 1 degree or smaller cells, with a scale from
1–5 from low to high diversity. The actual values given in the legends are dependent on the total
number of taxa in the area. A third map in this section depicts ‘rarity’ that is it distinguishes cells
with species that occur only there or in few other cells from those that are more widespread and
gives the cells a rating on a scale of five values. Effectively these rarity maps depict the distribu-
tion of narrow endemics.
Photographs of conifers in the wild are included in many places, selectively and when avail-
able. These are captioned with species name and location, and may be accompanied by further
comment given in a text box. The emphasis is on habitat shots to give an illustration of the
environment a species occurs in; in many cases this can be linked to the map, which is a Natural
Earth relief map, and to comments on habitat in the text. An attempt has been made to give
examples of the wide variety of habitats in which conifers grow and to spread these evenly in the
chapters. Limited availability of good photographs from some regions is the reason why this has
not always been achieved.
The Conifer Database, with 36,755 records, is the source of this Atlas. We have made use of
these data in a limited context, namely to produce maps that show the geographical distribution
of the conifers. We believe we have exploited the data for this limited purpose to the full in this
Atlas. However, these data can be used in different ways, in a taxonomic context, but also for
wider biogeographical research. In the chapter Examples of further research we present in several
short paragraphs some examples, a few of which have been contributed by other researchers who
have used the Conifer Database. None of these examples are in any way research papers; they are
only pointing at possibilities for further analysis of the data.
Some further statistics are presented at the end of the Atlas in Tables and Indices. These sec-
tions could also easily have been expanded to derive more information from the basic data. As
with the examples in the final chapter, what we present here should be seen as possibilities aris-
ing from the Conifer Database, which is made available online (see chapter Examples of further
research; The Conifer Database online). We present statistical data in Table 10 by country on the
total number of taxa, endemism and conservation status in the IUCN Red List. In the Species
Index we give the number of degree squares in which a species was collected and the conserva-
tion status of each species in the IUCN Red List of Threatened Species™ as of July 2013.
This Atlas is the first publication to present maps of all extant families, genera and species of
conifers. Numerous publications exist with distribution maps of some kind, usually range maps
that shade an area, sometimes with dot maps indicating localities, but all are confined to a taxon,
a country or region, a taxonomic rank such as genus (e.g. Eckenwalder, 2009) or another subset of
all conifers (e.g. Debreczy & Rácz, 2011). Published maps of species in the temperate zones of the
Northern Hemisphere far outnumber those representing species in the Southern Hemisphere and
in the Tropics. For several species this Atlas presents the first published distribution map.
 Acknowledgements

The work on this Atlas has a fairly prolonged ‘prehistory’ because the maps are based on a
database of herbarium specimens that has been nearly 30 years in the making. Many botanical
institutions and persons were variously involved with this, but we will here limit the explicit
acknowledgements and thanks to those involved from the time the Atlas of the World’s Conifers
became a collaborative project between two researchers, Aljos Farjon and Denis Filer. We worked
at different aspects more or less simultaneously and we acknowledge here those organisations
and individuals under the appropriate headings.

Funding

Funds were required for certain aspects of the work, mostly data gathering and associated travel,
and we are very grateful for generous support given by Lawrence Banks, Barry Denyer-Green,
Terence Devonport, Francine von Finck, Robin Jamie, Arabella Killander and Richard Storey to
the Conifer Atlas Project. Their confidence and trust in the timely delivery of the goods is highly
appreciated.
Willard Hunnewell, Wellesley, Massachusetts, USA is thanked for generous support towards a
visit to the Harvard Herbaria (A, GH) and the Hunnewell Arboretum by the first author (AF).
The Kew Guild, Royal Botanic Gardens, Kew, is thanked for financial support towards travel
costs involving visits to the USA, the Netherlands and South America by the first author (AF).

Institutional Support

The Herbarium, Library, Arts & Archives department of the Royal Botanic Gardens, Kew is
thanked for general support in kind and access to facilities given during this project to the first
author (AF).
The Department of Plant Sciences, University of Oxford, is thanked for continuing support to
the second author (DF) and with the BRAHMS project.

Data Gathering

Our thanks go to all the curators of institutional herbaria around the world who have given access
to herbarium specimens over many years of research on conifers by the first author (AF). Her-
baria mentioned in the text are abbreviated according to Index Herbariorum (http://sciweb.nybg.
org/science2/IndexHerbariorum.asp).
The Allan Herbarium Specimen Database (Database Manager Dr. Aaron Wilton), Landcare
Research, New Zealand is acknowledged as the provider of herbarium specimen data from New
Zealand herbaria (AK, CHR, WELT).
Gennady Firsov and Larissa Orlova, Komarov Institute, Russian Academy of Sciences, St. Peters-
burg, are thanked for help with herbarium labels in Russian at LE.
Olga Korniyenko, M. G. Kholodny Institute of Botany, National Academy of Sciences of the
Ukraine, Kiev, is thanked for help with herbarium labels in Russian at KW.
Qiaoping Xiang, Institute of Botany, Chinese Academy of Sciences, Beijing is thanked for help
with herbarium labels and locality data in Chinese at PE.
xii acknowledgements

Nodoka Nakamura, Department of Plant Sciences, University of Oxford (in 2011) is thanked for
help with data from Japanese herbaria.
Michael Nee, New York Botanical Garden, New York provided collection data on conifers in
Bolivia.
Tim Waters, South Tottenham, London (formerly of Department of Plant Sciences, University
of Oxford), is thanked for herbarium specimen data relating to the genus Agathis.
Jenny Wakefield, volunteer at the Royal Botanic Gardens, Kew, is thanked for invaluable help
with data entry relating to Kew’s herbarium specimens.

Providing Other Information and Help

Justin Moat, HLAA GIS Unit, Royal Botanic Gardens, Kew, is thanked for help with base maps.
Jan Wieringa, National Herbarium of the Netherlands (Naturalis), Leiden, is thanked for help
with rarity calculation procedures.
Andrew Liddell, Department of Plant Sciences, University of Oxford is thanked for his work on
the development of the BRAHMS online service.
Various people are thanked for contributing photographs, all of whom are credited in the cap-
tions to the photographs [non-credited photos are by the first author AF].
Sarah Mathews, Arnold Arboretum, Harvard University, Cambridge, Massachusetts, is thanked
for sharing results of phylogenetic research and discussions.
David Frodin, HLAA, Royal Botanic Gardens, Kew, is thanked for discussions on biodiversity,
particularly relating to Malesia and Australasia and for help with finding pertinent literature.

Reviewing

Colin Hughes, Institut für Systematische Botanik, University of Zurich, Switzerland, is thanked for
comments on the Mexico, Central America and West Indies chapter.
William Hawthorne, Department of Plant Sciences, University of Oxford is thanked for com-
ments on the Africa chapter (the first to be drafted).
Jonathan Timberlake, HLAA, Royal Botanic Gardens, Kew is thanked for comments on the
Africa chapter.
Christopher Page, University of Exeter, Devon, UK is thanked for comments on the Malesia
chapter.
Tim Waters, South Tottenham, London (formerly of Department of Plant Sciences, University
of Oxford), is thanked for his invaluable work as copy-editor of the Atlas.

Publication

Michiel Thijssen (Brill) is thanked for help and support with the publication throughout its cre-
ation, including a flyer printed by Brill to promote the book during the project to potential donors
and others.
Sabine Steenbeek (Brill) is thanked for technical advice on various aspects of the publication.
 Introduction

The natural distribution of the 615 species of conifers recognized in A Handbook of the World’s
Conifers (Farjon, 2010a) is the subject of this book. Conifers are a monophyletic group of gymno-
spermous woody plants with a long evolutionary history dating back to the Carboniferous more
than 300 million years ago (Ma). The fossil record is relatively abundant throughout most of this
long time span and it shows that conifers spread to a world-wide distribution by the beginning of
the Mesozoic ca. 250 Ma and attained their greatest diversity during that era (Stewart & Rothwell,
1993). For much of this time conifers were with few exceptions the only forest forming trees until
angiosperms appeared during the Early Cretaceous (Aptian) ca. 120 Ma. Competition with these
newcomers from the Late Cretaceous onward has resulted in exclusion of conifers in certain
areas or habitats. They are absent in the Amazon and Congo Basins and absent or rare in the
lowland plains of NW Europe and in the steppes of Asia, North America; South America. Conifers
no longer occupy salt marshes and only occur behind mangrove zones out of reach of frequent
incursions of the sea. There is evidence that they occupied such salty habitats during the Jurassic
and Cretaceous, with apparently truly halophytic species in the extinct family Cheirolepidiaceae
(Watson in Beck, ed. 1988). Despite these exclusions, conifers are still cosmopolitan and occur on
all continents except Antarctica (which no longer has any woody plants). They are present in all
climate zones from the Arctic and Subantarctic to the Tropics, but absent at the highest latitudes
where plant growth is severely inhibited. The same is true in extreme deserts, where higher plants
are mostly absent.
As with all classes of organisms, the diversity of species of conifers increases with decreasing
latitude, so generally there are more species per unit area at lower latitudes. However, this gen-
eral trend is distorted by the configuration of land masses and islands, the position of mountain
ranges and the position of unsuitable habitat, so that species are unevenly distributed across and
within climatic zones and greatest diversity of conifers is reached at different latitudes N and S
of the Equator. Dispersal capacity and history, speciation by isolation as a result of dispersal or
vicariance, and extinction are other factors that helped determine the natural distribution of
conifers. All these factors influencing the distribution of species have caused an extraordinarily
diverse pattern. While some species in the same genus are confined to extremely small areas,
others spread across a continent or, in one case, even wider. Species which are not closely related
can have very similar distributions, not only in their extent of occurrence (range size) but also by
largely overlapping (sympatry), often not only horizontally (extent) but also vertically (altitudinal
range). Similar habitat preferences of such species are but one possible explanation, today often
used to predict occurrences in areas from where no records exist. However, all distributions have
a history of which the present situation is likely to be a ‘time capsule’ instead of stasis indicative
of the past or the future. The very different distributions of closely related species may indicate
differences in habitat preference, or they may be the result of historical events. Extinction often
appears to have been an important factor in determining range sizes of species, as evidenced by
the fossil record in several well documented cases. Perhaps we are witnessing this process at work
in the differential demise of two related pines with similar habitats and mostly sympatric ranges
in W North America. Pinus albicaulis is declining rapidly in parts of its wide range due to attacks
by two different pathogenic organisms; Pinus flexilis is not affected. The retreat into refugia of two
European spruces during the glacial expansions of the Pleistocene epoch and their subsequent
spread in warmer times is another example of this historical factor in present distributions. Picea
abies returned to occupy much of its former range each time, albeit successively less far into
2 introduction

Atlantic Western Europe; Picea omorika was much less successful and is now confined to one
small refugium in the Balkans. When planted in other parts of Europe P. omorika does very well
and can often spread by itself from these plantations. The latter species is closely related to Picea
mariana in North America, which has one of the widest distributions of all species in the genus.
Comparative mapping, implicit in the Atlas by presenting all distributions of families, gen-
era and species in one volume and explicit in several additional maps, can reveal distribution
patterns more clearly. Several conifer genera, for instance, display similar disjunct distributions
on either side of oceans, in the Northern Hemisphere as well as in the Southern Hemisphere.
There are significant correlations with plate tectonics and the long history of continental drift,
but extinctions also played an important role leading to present patterns of distribution. Another
factor of importance is dispersal. Long distance dispersal, once virtual anathema in cladistic bio-
geography, is increasingly invoked in connection with the more recently emerged ability to esti-
mate divergence time in molecular phylogenetic analysis. This is mainly due to the relatively
recent dates obtained for much of the divergence into extant species within ‘crown clades’ of the
resultant hypothetical phylogenies. Often the separation of the relevant land masses resulting in
intervening oceans is much older than the divergence between the taxa they support. Evidence
from ecology, palaeobotany, geology and phylogeny reconstruction has to be evaluated in com-
bination in order to understand these intriguing patterns of conifer distribution.
With cumulative mapping, i.e. adding records of multiple species on to one map, we have
demonstrated several distribution patterns that are of great interest biogeographically. As often
as not, these patterns are as much revealed by absence as by presence, in fact they are of course
the result of both. These distribution patterns show connecting ‘corridors’ such as the Central
American isthmus and the Himalayan range, as well as gaps such as the one in the Andes between
30° S and 33° S and another between the Caspian Sea and the Hindu Kush. The gap in the Andes
has never been published for conifers before and its demonstration is the unambiguous result of
our strictly applied methods of verifiable data gathering and use.
The Swedish palaeobotanist Rudolf Florin published the first global atlas at family and genus
levels of all known conifers, both fossil and recent (Florin, 1963). The present Atlas of the World’s
Conifers is much more inclusive and detailed both because of its inclusion of all species in addi-
tion to these higher ranks and due to its methods, using techniques not available to Florin. How-
ever, it excludes maps of the fossil record of conifers. Florin understood clearly the importance
of that record for the interpretation of present distributions. The fossil record is very incomplete
and despite continuing new discoveries it will always remain incomplete. This is not a reason to
neglect it and replace it with more hypothetical interpretations. It is reason to intensify research
on all fronts, for which this Atlas is intended as a source of information.
 Creating the Atlas of the World’s Conifers

Introduction

This chapter discusses technical aspects of creating the Atlas. We include details about how data
were gathered and assembled into a database; the sources of data; the pitfalls one can encounter
especially when working with virtual data; and how the various categories of maps were pro-
duced and limitations with their interpretation. We also comment on the value of the database
as a whole to a project such as this. The data underlying the Atlas are available online providing
a means to verify the distribution of all conifer taxa. The data also constitute a valuable resource
for conifer researchers and those interested in the bigger picture of plant biogeography.

Data Gathering

The Atlas of the World’s Conifers is based on information obtained from herbarium specimens
preserved in institutional herbaria. Each collection, which may be a unicate or several specimens
as duplicates in several herbaria, constitutes a record in the Conifer Database. This database
contains some 37,000 records from 337 herbaria. The database was initially developed as an infor-
mation source for taxonomic work on conifers, not with the view to create distribution maps of
all taxa. There was an emphasis on monographic work in certain families but not in others and
subsequently a representative sampling of all taxa was added for work on the Handbook of the
World’s Conifers (Farjon, 2010a). Although the database had grown to a substantial size over the
years, it had to be expanded for nearly all taxa if maps based on herbarium collection localities
were to show reliable patterns of distribution.
Many older herbarium specimens do not have detailed locality information and are useless for
a distribution map. The data entry for the Atlas had to be focussed on identification and locality.
A renewed effort was needed, involving comprehensive data gathering in several major herbaria,
in particular the Herbarium of the Royal Botanic Gardens, Kew (K, the first author’s home insti-
tute). It soon became apparent that the examination of real specimens by a single researcher,

Table 1. Herbaria with >1000 records in the Conifer Database. [Notes: collections can have duplicates in
several herbaria. Taxa are species + infra-specific ranks.]
Herbarium Acronym Records Taxa Countries
RBG Kew K 9382 730 142
RBG Edinburgh E 2373 459  94
Missouri Botanical Garden MO 2201 422  85
New York Botanical NY 2006 289  47
Garden
Arnold Arboretum A 1743 272  50
Herbarium CAS Beijing PE 1644 195  14
Stockholm S 1631 354  99
NHN Leiden L 1538 161  58
MNHN Paris P 1488 230  52
NHM London BM 1221 266  76
4 creating the atlas of the world’s conifers

even when assisted by volunteers, could not produce enough data within a reasonable time. Data
sources available through electronic media had to be considered. These were used to various
extent, depending on the detail they contained and their estimated reliability. No datasets or
sections of these were ever imported in the Conifer Database wholesale, instead each record was
scrutinized and if found accurate it was entered individually.
The production of a map that claims to represent the distribution of a taxon, even when draw-
ing on virtual sources of data and using the latest wizardry in mapping software, involves hard
work. There are few shortcuts when creating reliable distribution maps.

Strategy for Data Gathering: Exclusions and Data Quantity

The Atlas presents the natural distributions of taxa, from family rank down to variety, of the coni-
fers of the world. This obviously excludes occurrences of planted conifers in gardens, parks, as
roadside trees or in commercial forestry plantations. Similarly, introductions to other continents,
even if naturalized, are excluded (e.g. the genus Pinus in Australia or Pseudotsuga in Europe).
However, there are situations where the distinction between naturalized outside the native range
and truly indigenous has become difficult or even controversial. These difficulties are invariably
linked with an ancient history of tree planting; the distinction between indigenous and intro-
duced in the case of conifers is clear in a country such as New Zealand. Europe and China are the
main regions where these problems arise, but more documentation is available in Europe, where
we can date most introductions of trees. Even if a species is indigenous in China that does not
mean that where we find it now it occurs naturally. In the absence of documentation about plant-
ings, ecological criteria may be used: if a species naturally occurs in primary forest but records
are from villages with surrounding secondary vegetation and disturbed habitat, we may have
reasons for doubt. For some species the reconstruction of the natural range may have become
so difficult that we have to adopt a more permissive approach or exclude the species altogether
from a map. An example is Pinus pinea in the Mediterranean (map EM-10 on p. 147), planted at
least since Roman times for its edible seeds if not for its ornamental values. In Europe, naturalized
introductions from before 1500 CE are usually included in Floras as native because they may date
from prehistoric times and came in slowly with the practice of agriculture. After 1500, voyages of
exploration began to bring back exotics from other continents and we often know which species
these are. Much of this also happened in E Asia, where Ginkgo biloba was introduced to Korea
and Japan from China many centuries ago, while other trees travelled in the opposite direction,
e.g. Cryptomeria japonica (Farjon, 1999). The documentation of this is scarce or non-existent, at
least in a scientifically verifiable form. Working from herbarium specimens with often scant or
even absent locality or habitat notes, the decision to include or exclude for the Atlas had to be
made on circumstantial evidence and ultimately on judgement derived from expert knowledge.
For example, in much of China, Platycladus orientalis is considered to have both medicinal and
spiritual value so it has been planted all over and often naturalized, as far away as the Elburz
Mountains in N Iran. Natural forests with this species as a component are very rare and probably
limited to a few provinces in N-central China, to where we have limited its natural occurrence
(map MAJ-107 on p. 241).
How many dots do we need on a map to represent the natural distribution of a species? This
depends on the range of the species, the topography within that range, the actual area of occu-
pancy (AOO), and the overall distribution of individuals, which may be contiguous, scattered, or
widely disjunct. Species with a wide range require more map points than narrowly distributed
species; species in an archipelago require fewer dots than those spread on continents, unless
there are intervening areas as unsuitable as the ocean. Some species are forest forming trees,
others occur sporadically or in special habitats which are unevenly distributed. The overall
aim is to present the extent of occurrence (EOO) and patterns of distribution within that range.
Data availability varies widely in the herbaria, from a single specimen to several hundred per spe-
cies. The latter is merely a rough estimate, because no effort has been made to gather all the data
 creating the atlas of the world’s conifers 5

for these abundantly collected species. Instead, selective data gathering has to be the strategy
for reasons of quality control, resolution and a more or less even distribution of map points, low
precision of geographical data (to the nearest minute at best) and available time for the task. The
point of diminishing returns can be reached sooner than anticipated. For several very widespread
species herbarium specimen data would remain insufficient even if all available material could
be included. This is particularly the case with the conifers of the boreal forest zone (taiga forest),
which not only have very large EOOs but are ubiquitous and undercollected within these ranges.
As we have restricted map points to those linked to herbarium specimens, the maps for these
species remain very incomplete and this has been pointed out where appropriate.

Strategy for Data Gathering: Data Quality

Potential sources of map data include field observations, herbarium specimens and published
localities other than herbarium specimen citations. We have chosen to use data gathered from
herbarium specimens only. The main reason for this is that verification is possible; there is at
least one specimen preserved in an institutional herbarium with access to the general public
for every dot on the map. The data used for mapping in the Conifer Database are accessible on
a website (see chapter Analysis and further research) including tools to associate dots on maps
with information in the database. The sources of herbarium data are varied and, apart from the
actual specimens lodged in herbaria, have greatly increased in this electronic age. There is no
doubt that the actual specimens remain the preferable sources of data for reasons of taxonomic
identification. Another reliable source is the specimen citations in monographs and taxonomic
revisions, where a specialist has done the identification. We will here concentrate on the various
sources of herbarium specimen data available in electronic form. These can be listed in order of
reliability for mapping purposes.

A) High resolution images of specimens that can be accessed by others, normally linked to online
virtual herbaria.
B) Data from well curated herbaria visited online or requested and sent as electronic files.
C) Data from lesser known herbaria visited online.

As we move from A to C, increasing care is needed to verify the data before they can be incor-
porated in the database. For B and C there are no images of specimens and we have to gauge
our confidence in the identifications indirectly. There are various criteria that may apply. Good
herbarium curation ensures that the naming of specimens is regularly updated and that this is
reflected in ongoing updates of the (online) databases. Well known herbaria are more frequently
visited by specialists who can correct misidentifications than are many smaller, often regional
herbaria. The content and quality of their databases also vary; the best have included all avail-
able label information and may even have added map reference data to localities. Confidence in
identification increases when new identifications are signed, we then know who has had a look at
the specimen and if an acknowledged specialist, we may take it up. Confidence in identifications
is also directly related to taxonomic or geographic distinction; if we know that there is only one
species of Taxus native in Europe then specimens collected in Europe from yews growing in the
wild belong to that species, even though yews are notoriously difficult to identify. We can include
the data from online databases as long as localities fall within the known range and wild status is
verified or likely. The determination of duplicates of one and the same herbarium collection may
differ between specialists; they may even differ over time by the same specialist. In the Conifer
Database, these may thereby end up as two different species. In some cases where there is such
ambiguity we have maintained both records.
A few sources of electronic data in the first two categories may be briefly mentioned here as
examples. An example of category A is the Chinese Virtual Herbarium database (http://www
.cvh.org.cn) which gives access to herbarium specimen data of Chinese herbaria, many with
6 creating the atlas of the world’s conifers

images of specimens. This source has been used extensively for the Atlas, despite the difficul-
ties imposed by language (an online translator can be used as long as it gives pinyin transcrip-
tions of Mandarin characters besides ‘translations’). Many herbaria consulted electronically fall
under B, some of which did not give access online but sent requested data as Excel files (e.g. the
three main herbaria AK, CHR and WELT in New Zealand). Good examples of online sources are
the Consortium of California Herbaria (http://ucjeps.berkeley.edu/consortium/) and the Rocky
Mountain Herbarium (RM) in Laramie, Wyoming (http://www.rmh.uwyo.edu/) with extensive
data for California and the Rocky Mountain states in the USA. The CCH combines data for the
state’s flora from all institutional herbaria in California, RM is a herbarium that specializes in the
Rocky Mountain flora. Another consortium of herbaria databases exists for the Pacific North-
west states of the USA and British Columbia (http://www.pnwherbaria.org/). Many herbaria
have pooled their specimen data in the Global Biodiversity Information Facility (GBIF) (http://
data.gbif.org/) and these can be accessed in combination. The search is in the case of the Atlas
facilitated by the fact that GBIF has adopted the taxonomy used in the Catalogue of Life (http://
www.catalogueoflife.org/) in which the Conifer Database has been compiled by the first author,
but alternative names are in many cases also given. GBIF gives access to a great variety of data
sources, including observational data, for the occurrences records of taxa and yet undertakes no
verification of these data; GBIF is a portal to these individual datasets. Selective use is paramount
to keep quality standards in the database high and in most cases only data from herbaria consid-
ered to belong to category B (images were not available via GBIF) have been considered. A few
standardized assumptions in GBIF can be false: ‘year 02’ is interpreted as 2002 but can be 1702,
1802, 1902 or 2002; altitude is interpreted as given in meters but can be feet. Parts of the data have
been given map point references in decimals or degrees; errors were encountered frequently and
so all had to be checked.

Geographic Data and Building a Gazetteer

The main categories of geographic data from all sources are broadly continent, country, major
country area, (state or province), minor country area (municipality or county), locality name
(small populated place, mountain, reserve), locality notes (“2 km west of Nadja Paka”), and a map
point reference in degrees and minutes. Although the objective is to get to the map point, enter-
ing higher level data is useful to produce different outputs and for data verification. Discerning
even these categories of data, especially in foreign languages but also on older herbarium labels,
can be difficult. Frequently not all categories are stated and they need to be added once the local-
ity has been found on a map or in a gazetteer. Until the advent and subsequent routine usage in
the field of hand-held Global Positioning System (GPS) devices, few collectors added map point
references in degrees and minutes to their label data. The search for localities in gazetteers and/
or on maps in order to add map point references is crucial, but it is also by far the most time
consuming element of the data gathering process. Generally, the difficulties increase with the age
of the specimen, even if a locality is mentioned, which often it is not. For instance, J. D. Hooker’s
herbarium collections at K from the Himalaya, made in 1849, often give “Sikkim” which is a large
area. Other pitfalls are changes in place names over time, or renderings of hearsay toponyms by
collectors using their imagination rather than a standardized system of transcription, e.g. from
Chinese to pinyin as is now common practice. Some collectors indulged in a route description
(“12 miles north along Hwy. 123 from the fire lookout, then 4 miles on the left hand dirt road”)
more as if wanting to give directions for a revisit than to pinpoint the locality on a map. For this
database, as it was built up over many years, many sources were consulted to locate herbarium
collections. Recently, the primary sources have been two online gazetteers, often used simul-
taneously: JRC Fuzzy Gazetteer (http://dma.jrc.it/services/fuzzyg/) and GeoNames (http://www.
geonames.org/). If no locality name was given but it is possible to locate the collection to a map
point, e.g. with a given GPS reading, Google Earth can be used to find the nearest one for the
database gazetteer. From the herbarium specimens, a database gazetteer is compiled to which
 creating the atlas of the world’s conifers 7

Table 2. Salient features of BRAHMS relevant to the Conifer Atlas project.

Data capture Flexible and secure data capture optimized for speed and accuracy.
Species and geographic data verified on entry.
Taxonomic framework Facility to develop a central list of all taxonomic names, easily
sorted, located and organized into groups as required. Storage of
details for names such as distribution summaries, conservation
status, synonymy and common names.
Botanical records and specimens Separation in the database structure of individual collection events
(who, what, where, when) with the physical specimens as held in
herbaria.
Query tools Selection of data from main database into query files for further
processing. Typical queries used by the Atlas project to generate
maps, summaries and statistics are by taxa (species, genus, family)
and geographic unit (country, province, grid cell).
Mapping Generation of decimalized map points from BRAHMS query files
passed to a selected GIS and plotted.
Summarizing and indexing Creating custom summaries, statistics and indices, for example,
total specimens per herbarium, list of species per country, index of
botanical names with add-on statistics.
Diversity analysis The ability to calculate for any given area (e.g. country or grid cell)
total number of families, genera and species, collection density,
rarity and other diversity-related parameters.

individual herbarium records can be linked. The larger this becomes, the more it can be used to
locate other collections, as plant collectors often visit the same sites for obvious reasons. Besides a
gazetteer location in degrees and minutes N/S or E/W, a refined map reference may be necessary
to accommodate statements like “8 km NW of village”, which will become the map point. Map
data in the Conifer Database are rounded to the nearest minute.

Building a Database
For anyone embarking on a project that uses data gathered from a large number of specimens to
create maps and potentially other outputs, gathering data electronically is necessary. The choice
of software is down to individual preference and may range from a simple spreadsheet to a fully
relational database. While some computer literate botanists are capable of developing their own
data management systems, others will use something off the shelf and ready to go.
This Atlas has been created from BRAHMS (Filer, 2012), partly because the Conifer Database was
developed in this system from the outset but also because BRAHMS provides data processing and
analysis features relevant to a project such as this. Some of these key features are listed in Table 2.

Mapping

Mapping Software
All maps in the Atlas were created by selecting the relevant data points from the BRAHMS data-
base and transferring these to a Geographic Information System (GIS). As with the database itself,
the choice of GIS is largely down to familiarity and user preference.
We elected to use DIVA-GIS for almost all maps presented in the Atlas. DIVA-GIS is a free
computer programme for mapping and analysing data, such as the distribution of species or other
point-distributions. DIVA-GIS reads and writes standard data formats such as ESRI shape files.
BRAHMS communicates directly to DIVA-GIS facilitating map production. The salient features
in DIVA-GIS used to produce the maps are given in Table 3.
8 creating the atlas of the world’s conifers

Table 3. Salient features of DIVA-GIS relevant to the Conifer Atlas project.

Freely available, well
documented, easily installed
Map layers
Importing data points
Point and polygon attributes
and data classification
Diversity analysis
Export map images
Can be downloaded for free from http://www.diva-gis.org/ together with a
useful technical guide.
Uses standard ERSI vector shape files (SHP) for map layers, a vast number
of which are freely available online. It can also read raster files such as
the TIF files used for most of the Atlas base maps.
Imports data coordinates from various sources, including DBF, Excel and
text file formats.
Data point symbol, colour and size are easily controlled and made unique
on any selected field (e.g. points on multiple species maps differently
coloured); polygon areas can be classified, shaded, etc.
Useful spatial analysis features to calculate and display species richness
by grid cell area using colours.
Map images easily saved in high resolution for printing.

Basemaps and Mapping Points

Most of the Atlas maps use the Natural Earth raster relief base maps as available on http://www
.naturalearthdata.com/. Here, one can access free vector and raster map data at 1:10,000,000,
1:50,000,000, and 1:10,000,000 scales. From the selection provided, we chose the large scale data,
1:10,000,000 raster Natural Earth II base map. Depending on the continent or map region, further
vector layers were added (country and in some cases administrative boundaries). For smaller
regions and islands, the Natural Earth II base map had insufficient resolution and was replaced
by a plain vector map or if available by another type of relief map.
Map points have mostly been plotted as black dots. In some cases, we have used red dots to
provide contrast with a complex base map background (for example, some maps in southern
Chile and parts of the Himalaya). Where maps have two or more taxa, these are mostly differenti-
ated by symbol colour.

Interpreting the Maps

In the Atlas, a dot on the map (map point) represents the locality where a herbarium collection
(or collections) was made. Map points are distributed first and foremost according to the avail-
ability of herbarium collections with locality information in the consulted herbaria. In all maps,
the objective has been to present an overall distribution as near to the actual distribution as the
available data allow. However, if a species is known to occur in a certain area but no specimen
was available, there will be no map point in that area. When there were large numbers of her-
barium specimens of a species in many herbaria, the data were entered selectively to obtain the
overall distribution, not the greatest possible density of map points. Special efforts were made to
map outlying occurrences wherever these were known and specimens had been collected there.
The extent of occurrence (EOO) or range of a species determines the scale of the map and this
influences the density of map points per unit area, independent of data availability. The smaller
the EOO, the greater the density of map points required to give a reliable impression of distri-
bution. The greater the EOO, the more difficult it is to find enough herbarium records to cover
it; there is also the law of diminishing returns in these cases. For ubiquitous species that occur
across continents, e.g. in the boreal forests of Canada and Russia, herbarium collections are inad-
equate even if all existing specimens could have been entered in the database. For Canada few
and for Russia almost none of these are available in electronic databases. Could we have used a
different approach to map these species? Raster maps that shade the regions where these species
occur exist, but their accuracy varies greatly. Instead, we have indicated the boreal forest zone
 creating the atlas of the world’s conifers 9

on several maps and explained the inadequacy of the map points to represent the distribution
of these taxa.
Maps of genera and families in the Atlas combine the map points in the database for two or
more species. The density of map points per unit area is thereby inevitably increased; the quan-
tity of dots per unit area is proportional to the number of species in the area. Therefore, maps
of genera and families give a measure of species diversity in areas with greater density of map
points and not of abundance. In general, there may be no necessary relation between density of
map points and the abundance of conifers.
Diversity and rarity maps are given for ‘continents’ and sometimes smaller areas. The legends
in these maps are relative to the area of each individual map only and do not compare with other
maps. In all we have created a five-scale legend using different colours, with green as the lowest
and red as the highest value. In the diversity maps the scale represents numbers of taxa in five
classes, but in the rarity maps the values are scores on a scale that have no intrinsic meaning and
these are therefore omitted. These maps, although they look similar, present two different aspects
of conifer distribution. The diversity maps highlight areas where greater numbers of genera or
species occur; they point at ‘hot spots’ of conifer diversity. The rarity maps give the distribution
of narrow endemics, species which only occur in a very limited area. Often the narrow endemics
occur in areas with higher diversity, but there are exceptions and comparison of the two catego-
ries of map may reveal these.
Three example websites used to locate and geo-reference herbarium collections for the Atlas of the World’s Conifers.
Data from all online sources were used cautiously and only when the identification and locality details were trusted.

The Chinese Virtual Herbarium http://www.cvh.org.cn/data/contains many imaged specimens (red symbol in left hand
image). These can be viewed together with the data summary. Headings can be viewed in English but content is in Mandarin
characters; this has to be put to an online translator for non-Chinese users. Online gazetteers can assist with finding
localities. Most of the geo-references given with the specimens are only at county level and/or need verification.

The Rocky Mountain

Herbarium (RM),
University of Wyoming,
http://www.rmh.uwyo.edu
/data/search.php, showing
specimen search results for
a species in List and Detail
mode.

Global Biodiversity Information Facility (GBIF) http://data.gbif.org/occurrences/. Mapping in GBIF datasets has been
accomplished in only a percentage of specimens, depending on the herbarium source. The geo-references require verification.
 Global and trans-continental distributions

In this chapter we present a world map with the distribution of all conifer species and maps of
the families and genera with a wide, trans-continental but not highly fragmented distribution.
Small maps of all families are given together to compare their global ranges, but those with a
more limited distribution are presented and discussed in the following chapters relevant to where
they occur.

Global Distribution of Conifers

Map GTC-1. Global distribution of all conifer species. The boreal conifer forests of Alaska, Canada, Scandinavia
and Russia are shaded in this map to show their extent. Available localities from herbarium collections are
inadequate to show the extent of occurrence (EOO) and area of occupancy (AOO) of the widespread boreal
conifer species. [Source of data on extent of boreal forest zone: www.fao.org/geonetwork/srv/en/metadata
.show?id=1255]

The global distribution of all 615 conifer species shows distinct patterns. In the N a belt of boreal
coniferous forest dominated by a small number of species encircles the globe; it is shaded on the
map and would be dense with dots if the herbarium collection data had been available. To the S of
this boreal forest zone the nearly 37,000 herbarium specimen records in the Conifer Database give
an accurate general picture of conifer distribution at this scale. One pattern that stands out is that
conifer species are most abundant in major mountain systems, e.g. in W North America, Mexico
and Central America, in the Andes of S Chile, in Europe from Spain to Greece, in the Himalayas,
China and Japan, and in New Guinea and New Zealand. Other areas of abundance are perhaps less
expected, such as the Atlantic side of the USA and SE China, where forests tend to be dominated
by angiosperms. Less densely populated but equally surprising may seem parts of Australia and
parts of Malesia other than New Guinea, where angiosperms are once again the dominant trees
of the forests. Some small islands are rich in conifers, most notably New Caledonia, Taiwan and
Tasmania. Where are conifers scarce or absent? Apart from regions such as Antarctica, the High
Arctic and major tundra zones, steppes and deserts, there are some very large areas devoid of
conifers where this absence is perhaps less obvious. In South America these are the Amazon Basin
and Mato Grosso, in Africa the Congo Basin and West Africa, in Asia the Indian subcontinent and
most of the Tibetan Plateau. For the latter, adverse climate is probably the cause, with a growing
12 global and trans-continental distributions

season too short for shrubs and trees due to its very high altitude. The Amazon and Congo Basins
are excellent habitats for tree growth; they harbour the world’s largest tropical rainforests. Coni-
fers occur in tropical rainforests, mostly in uplands but in some regions also in lowlands. In the
latter, they tend to occupy sites with very poor soils, where tree growth is much less vigorous and
consequently competition less intense, leaving space for slower growing conifers. Are such sites
absent from the Amazon and the Congo basins? The rivers that come into the Amazon from the
N, especially the Rio Negro, carry water very low in minerals, indicating that they spring from and
run through regions with poor soils. Ecology seems not a sufficient explanation and possibly the
absence of conifers in these large tropical river basins also has historical (contingent) causes. The
absence of conifers in West Africa is even more of a conundrum, perhaps until it is realized that
the entire sub-Saharan continent is poor in conifer species (see Africa chapter for more details).
Much of Africa has been subject to major and rapid climatic fluctuations during the later Neogene
which will have caused the extinction of many forest species (website: natureNEWS (4/1/2011): The
Drying of East Africa). This must have made it less likely for the remaining species to be able to re-
colonize; indeed only one species, Podocarpus milanjianus is now extremely widespread in Africa
(map AF-18 on p. 469) and it is inferred to have been the species that spread during periods of
global cooling in the Pleistocene (Morley in Turner & Cernusak, eds., 2011). This leaves the great-
est enigma of all: the near absence of conifers in the Indian subcontinent. All but one species of
conifer in India are confined to the Himalaya or to the hills in the far NE of the country and these
nearly all belong to Pinaceae, the subfamily Cupressoideae in Cupressaceae, or Taxaceae and are
of Laurasian origin (see below). The exception is Nageia wallichiana (map MA-55 on p. 323) with
a disjunct population in the Western Ghats of S India. During an extended period in the Meso-
zoic and early Cenozoic, India was a continental craton moving on the Indo-Australian Plate as
a large island northwards, from temperate S latitudes through the tropics to tropical/temperate
N latitudes and finally colliding with the Asian continent, pushing up the Himalaya and adjacent
mountain systems. India has a good Mesozoic record of fossil conifers, all of Gondwanan affinities
(Sitholey, 1963). Conifers went extinct during that long journey, but what caused this? Perhaps
climate change or the disaster of the Latest Cretaceous Deccan Traps, an event that itself caused
climate change, or a combination of these factors. In a subcontinent with elevations generally
below 1000 m and situated for the most part in the tropics, the natural vegetation is tropical to
subtropical forest dominated by angiosperms. India could only have been repopulated by conifers
from Asia, and few apparently succeeded to gain a foothold. Of these, only Nageia wallichiana, a
broad-leaved tree species in the Podocarpaceae, survived into the present, occupying a few loca-
tions in the Western Ghats mountains which rise to >2500 m.

Global Generic and Specific Hotspots

Not only are conifers unevenly distributed across the world (map GTC-1 on p. 11), there are also
disparities in the taxonomic diversity among the regions in which they occur. Some areas are
diverse and others are poor in genera and/or species. To show where high diversity is concen-
trated we have chosen to calculate the diversity of conifers at the spatial level of 1 degree cells
and plot the highest diversity levels only on global maps.
All eight families, 83% of 70 genera, and slightly over half of all conifer species occur in 14 centres
of diversity around the Pacific Ocean (Farjon, 2008). First noted by Engler (1926), Li (1953) sought
to explain this concentration around the Pacific at a time before continental drift and plate tec-
tonics were better understood and accepted. The centres of conifer diversity around the Pacific
Rim are all mountainous, often on islands, and have oceanic climates with moderate tempera-
tures and high precipitation. Separation and isolation, especially on islands, promotes specia-
tion. The great age of the Pacific as an ocean basin, going back to the origin of the oldest extant
families, also played a role. Continuous movement of its plates towards shifting land masses,
creating subduction zones where they contacted and thereby causing volcanism, island-forming
and mountain-building, provided suitable habitats during the entire period of the evolution of
 global and trans-continental distributions 13

Map GTC-2. The distribution of degree cells with ≥ 6 genera per cell shows the areas and hotspots of
generic diversity in conifers.

Map GTC-3. The distribution of degree cells with ≥ 10 taxa per cell shows the areas and hotspots of diversity
at species and lower ranks.

Map GTC-4. The distribution of degree cells with highest rarity values shows the distribution of conifer
taxa with very small ranges (narrow endemics).
14 global and trans-continental distributions

modern conifers. Other major centres of conifer diversity are in China (especially Sichuan and
Yunnan Provinces) and along the Himalayan Range from the E, with species diversity receding
further W but generic diversity remaining high. On the Atlantic Ocean coasts, only the NE of
the USA scores high in generic diversity but not in species numbers. Scattered degree cells with
high diversity are also present in W Malesia (Sundaland) more distant from the Pacific Rim, and
in Europe. The diversity in Europe is a remnant of much greater diversity before the Pleistocene
extinctions that devastated the tree flora of that continent.
Map GTC-4 shows the global distribution of narrow endemics. There are similarities and differ-
ences between the two diversity maps and this map that are notable. The large region with high
diversity in E Asia, Malesia and Australasia coincides with the distribution of narrow endemics
except that the latter do not occur in New Zealand. Mainland Australia has only one degree cell
with high generic diversity but four with narrow endemics, all are on the E coast. The W coast of
North America down to Central America also shows similarities in the distribution of diversity
and endemism. However, while diversity is low in the West Indies, there are a number of very
restricted endemics and the same situation prevails in the Mediterranean. A few narrow endem-
ics also occur in Africa and Madagascar. Not unexpectedly, island endemism accounts for a large
proportion of the distribution of these localized taxa. In some instances, taxa are known only
from the type locality due to incomplete collecting or problems with identification and these
may turn out to be more widespread, but nearly all cells in map GTC-4 represent true endemic
species, subspecies or varieties.

Distribution of Families

Eight families are recognized in conifers (Farjon, 2001, 2010a) and both the number of taxa included
and their geographical distribution vary widely, from monospecific (Sciadopityaceae) to 231 spe-
cies (Pinaceae) and from a single country (Japan, Sciadopityaceae) to all continents except Ant-
arctica (Cupressaceae). It is of interest to compare the distributions at family rank here in a series
of small maps; five families with wide distributions are then presented and discussed in more
detail while three families with smaller ranges are treated in separate chapters.

The Cupressaceae is the only family

Map GTC-5. The global distribution of Cupressaceae.
with a cosmopolitan distribution,
i.e. present in both the Northern and
Southern Hemisphere and on all
continents except Antarctica. This
distribution in both hemispheres
was already established during the
Early Cretaceous. The present wide
distribution in the Northern Hemi-
sphere is largely accounted for by
the genus Juniperus.

The Pinaceae is the Northern Hemi-

Map GTC-6. The global distribution of Pinaceae.
sphere family with the widest dis-
tribution; only one species crosses
the Equator in Sumatera. The fossil
record since the Early Cretaceous
establishes this family as Laurasian,
i.e. with an origin in the Northern
Hemisphere supercontinent. The
wide distribution is shared by sev-
eral genera, but with Pinus making
the largest contribution.
 global and trans-continental distributions 15
Map GTC-7. The global distribution of Taxaceae.
Map GTC-8. The global distribution of Cephalotaxaceae.
Map GTC-9. The global distribution of Sciadopityaceae.
Map GTC-10. The global distribution of Podocarpaceae.
The Taxaceae is a family with a
mostly Northern Hemisphere dis-
tribution, but two species occur
south of the Equator in Male-
sia and Australasia. One is the
monospecific Austrotaxus with a
probable Gondwanan, i.e. South-
ern Hemisphere origin. The fossil
record dates back to the Jurassic
and is only known with certainty
from the Northern Hemisphere.
The Cephalotaxaceae is a Northern
Hemisphere family of one genus
with at present a limited distribu-
tion in E Asia. This sister group to
the Taxaceae has a fossil record
dating from the Jurassic to the
Recent. It was once distributed
across Eurasia and North Amer-
ica, but has never been found in
the Southern Hemisphere with
certainty.
The Sciadopityaceae is a family
of a single species, most closely
related to the Cup­ressaceae. The
fossil record of Sciadopitys dates
back to the Late Jurassic. It shows
the family to have been present in
Eurasia, possibly widespread from
Europe to Japan. Sciadopitys ver-
ticillata is now confined to Japan.
The Podocarpaceae is a primarily
Southern Hemisphere family with
a pan-tropical distribution and
extensions across the Equator as
far N as Mexico and Japan. The
fossil record, possibly dating back
to the Late Triassic, is Gondwa-
nan, i.e. the family is of Southern
Hemisphere origin. The genus
Podocarpus accounts for most of
the distribution of the family.
16 global and trans-continental distributions

The Araucariaceae is a family with

a disjunct distribution in the South-
ern Hemisphere, with an exten-
sion across the Equator in Malesia.
Although this distribution seems
Gondwanan, the fossil record, dat-
ing from the Jurassic to the Recent,
shows that the family was prob-
ably cosmopolitan during much
Map GTC-11. The global distribution of Araucariaceae. of the Mesozoic. Only three gen-
era make up the family, two with
extremely limited or fragmented
distributions.

The Phyllocladaceae is a Southern

Hemisphere family of one genus
with a limited distribution in Tas-
mania, New Zealand and Male-
sia, where the single widespread
species in that region crosses the
Equator. The fossil record goes back
to the Palaeocene in Patagonia
and Australia, indicating a Gond-
Map GTC-12. The global distribution of Phyllocladaceae. wanan origin possibly in the Late
Cretaceous.

The principal division in the distribution of conifer families is between the Northern and Southern
Hemispheres. In a phylogenetic context, extant conifers divide into (nearly) exclusively northern
clades (Pinaceae, early diverging Cupressaceae, subfamily Cupressoideae in Cupressaceae, Taxaceae
s.l., Sciadopityaceae) and southern clades (Araucariaceae, Podocarpaceae s.l., subfamily Callitroi-
deae in Cupressaceae) (Leslie et al., 2012). Although much of the taxonomic diversity within these
families is of relatively recent (Neogene) age, their more basal lineages date from the Mesozoic,
with Araucariaceae, Podocarpaceae, Taxaceae and Sciadopityaceae diverging by the beginning of
the Triassic and Cupressaceae and Pinaceae by the Late Jurassic to Early Cretaceous (Leslie et al.,
2012 Fig. 1). Therefore, the principal division in the global distribution of conifers is likely to be
related to the Laurasia/Gondwana separation of an earlier global landmass (Pangaea) which began
during the Triassic and was complete by the Late Triassic ca. 200 Ma. The presence in the fossil
record of Araucariaceae in both Laurasia and Gondwana since the Jurassic could indicate an origin
of this family predating the breakup of Pangaea, as suggested by molecular dating estimates.

The separation of Pangaea into Laurasia and Gondwana.

A taxon is always older than its oldest known fossils,

the more so when these are known from regions as
far apart as Argentina and England. Among the early
diverging Cupressaceae, of which the extant taxa as
well as the sister group (Sciadopityaceae) are limited
to the Northern Hemisphere, fossil remains dating
back to the Mesozoic (Triassic-Cretaceous) are known
from the Southern Hemisphere as well as the North-
ern Hemisphere (Stockey et al. in Farjon, 2005a). This
too indicates an origin dating from before the breakup.
 global and trans-continental distributions 17

The fossil presence in both hemispheres of the later evolving subfamily Callitroideae after the two
supercontinents had separated would imply one or more dispersal events. Much more recently
Africa reconnected with Eurasia, enabling Laurasian conifers to migrate; one species, Juniperus
procera, extending well S of the Equator. The family Podocarpaceae originated in the Southern
Hemisphere around the time of the breakup of Pangaea and remained there until much later
continents that had drifted apart during the Cretaceous were reconnected (or nearly reconnected
leaving only continental shelf seas open), enabling species to disperse northwards as far as Mexico
and Japan. The Pinaceae, originating in the Early Cretaceous (but possibly earlier) in Laurasia,
remained almost entirely in the Northern Hemisphere and just one species of Pinus spread S of
the Equator in Sumatera. This family was most successful in radiating more recently both in taxo-
nomic and geographic terms, largely due to the major differences in the distribution of oceans
and landmasses between the two hemispheres, with the greatest landmasses all situated N of the
Equator. The palaeogeographic history of Taxaceae is the least clear. Whereas its distribution is
almost exclusively in the Northern Hemisphere (with a dispersal case into Malesia S of the Equa-
tor involving one species Taxus wallichiana, see map MAJ-33 on p. 195), the monospecific genus
Austrotaxus is endemic to New Caledonia, which is a diversity hotspot of ‘southern’ conifers.
Together with vicariance events (separating populations by the creation of barriers such as
oceans) dispersal is a major cause of the present distribution of conifer species across the globe
(a third cause is extinction). In general, opportunities for long distance dispersal in the Southern
Hemisphere are limited in comparison to the Northern Hemisphere. Most conifers are ineffective
long distance dispersers; the exceptions are those which have evolved adaptations to dispersal by
birds. Among these, two genera in two families have been most successful in spreading widely,
Juniperus (Cupressaceae) in the Northern Hemisphere and Podocarpus (Podocarpaceae) in tropi-
cal latitudes. These two genera are largely responsible for the wide distributions of Cupressaceae
and Podocarpaceae, including their occurrence on oceanic islands. In Pinaceae, the genus Pinus
has been similarly successful even though only a few of its species are bird dispersed. With con-
tinuous or near continuous landmasses long distance dispersal is often not a necessary condition
for wide distribution, as long as suitable habitat is widely available. In this context it is notable
that the more diverse genera in the large families Cupressaceae, Pinaceae and Podocarpaceae all
radiated in the Late Cenozoic (Neogene) i.e. during the last 25 million years (My) (Leslie et al.,
2012). By this time the modern continents were not only formed, but had largely reached their
present positions on the globe. This is why species of conifers are much more numerous in the
Northern Hemisphere than in the Southern Hemisphere and why Podocarpus became much more
diverse in tropical latitudes than in the widely separated southern continents.

Cupressaceae

The family Cupressaceae is the third largest in conifers with 135 species but it has the widest
distribution. It is the only family with a presence on all habitable continents and is present on
many oceanic islands where conifers occur naturally.

Map GTC-13. The global distribution of Cupressaceae—detail.

18 global and trans-continental distributions

In the Northern Hemisphere N of the Tropic of Cancer (23° 26’ N) the distribution of Cupres-
saceae and of all conifers is similar. We have shaded the boreal coniferous forest as in map GTC-1
on p. 11 and for the same reason, but the only genus present here, Juniperus, is more scattered
in this zone than the various species of Pinaceae (map GTC-14 on p. 19). In the tropics the fam-
ily is much less widespread and occurs in New Caledonia, N Australia, New Guinea, Maluku,
S China (including Hainan and Taiwan), Indochina, the Arabian Peninsula, East Africa, the West
Indies, Central America and S Mexico. The family Cupressaceae is notably absent in Malesia W of
Maluku and in tropical South America. In the Southern Hemisphere S of the Tropic of Capricorn
(23° 26’ S) the family is present in New Zealand, Australia (including Tasmania), South Africa and
the southern parts of the Andes Range in Chile and Argentina. The partial division of the family
into two subfamilies Cupressoideae and Callitroideae is congruent with a division into Laurasian
and Gondwanan phylogeography (Mao et al., 2012) reflecting an middle Mesozoic origin and a
later Mezozoic radiation when the two super continents had separated. Incursions from Laurasia
to Gondwana and vice-versa may account for fossil and extant presence of basal Cupressaceae
taxa in Australasia and South America and fossil Widdringtonia in North America. In the tropics,
species of Cupressaceae are confined to high elevation locations, either above zones of evergreen
angiosperm forest or above a desert zone, as in the Arabian Peninsula. A few tropical species
can occur in the lowlands under adverse edaphic conditions, such as the ultramafic substrates of
New Caledonia and sandstone ‘breakaways’ in N Australia, but these appear to be exceptions. In
more southern latitudes under similar edaphic conditions species numbers can rise again, as in
Australia in the genus Callitris. Outside the tropics, species in this family have adapted to extreme
conditions of climate, so that we find particularly junipers ( Juniperus) under subarctic and even
arctic conditions N of the Arctic Circle as well as in alpine situations at lower latitudes, but also
in deserts. The only conifer species in the Sahara is Cupressus dupreziana and the Gobi Desert has
a variety of Juniperus sabina. Species adapted to moist or wet climates are mostly circum-Pacific
(Farjon, 2005a: 74) and several of these appear to have retreated there from wider distributions
in the past. Range contraction is evident in many of the extant species, especially those in the
early diverging clades of the phylogenetic tree (Leslie et al., 2012), whereas range expansion has
mostly occurred in more recently evolved clades, especially Juniperus but also Callitris in Austra-
lia. In both cases this has given rise to speciation and adaptation to various environments. As an
example of range contraction the palaeogeography of the subfamily Sequoioideae can be given.
At present, this group contains Metasequoia in E-central China and Sequoia and Sequoiadendron
in California. Fossils assigned to this subfamily dating from the Turanian-Santonian (Late Creta-
ceous) to the Pleistocene have been found across Eurasia and North America as well as in Aus-
tralia and New Zealand (Stockey et al. in Farjon, 2005a). The diversity as well as the geographical
extent of this group has been much greater than the three species that survived to the present.
If range contraction can be that great, the absence of the family Cupressaceae in large parts of
the world where conditions now are favourable must be seen as historical contingency. It had its
causes in changing environmental factors that influence distribution, but the particular events
responsible for extinction may be no longer traceable in the geological record.

Pinaceae

The family Pinaceae is the largest in conifers with 231 species (including some nothospecies, that
is, hybrids between other species) and although widely distributed, is virtually confined to the
Northern Hemisphere, with only Pinus merkusii crossing the Equator in Sumatera.
 global and trans-continental distributions 19

Map GTC-14. The global distribution of Pinaceae—detail.

The distribution of Pinaceae is similar to the distribution of all conifers N of the Tropic of Cancer,
reflecting the dominance of this family in conifer forests of the temperate to cold regions of the
Northern Hemisphere. Notable differences are found in semi-arid regions such as those in the
American southern prairie states, Iran and Afghanistan, where species of Cupressaceae occur
but those of Pinaceae are absent. In the tropics N of the Equator the differences are more pro-
nounced; there are no species of Pinaceae in South America (the southernmost occurrence is in
Nicaragua) nor in sub-Saharan Africa, nor the Arabian Peninsula, nor in the Indian subcontinent,
nor in Malesia except Sumatera and the Philippines. Pinaceae are dominant in the boreal forest
zone (shaded on Map GTC-14), with a limited number of species of Abies, Larix, Picea and Pinus
making up the taiga forest and woodland/tundra ecotone. Here the dots on the map would be
dense if enough herbarium collections had been available. To the S of the boreal coniferous forest
substantial gaps occur where steppe vegetation replaces the forest in the central part of the North
American continent and in various parts of Asia, notably around the Caspian Sea, in Kazakhstan
and Mongolia to Nei Monggol [Inner Mongolia]. In Europe, the family Pinaceae S of the boreal
forest in Scandinavia is present in most of the mountainous regions but absent in most of the
lowlands except around the Mediterranean and Black Seas and in the Baltic States, Belarus and
Russia. The Hungarian Plain stands out clearly as an open space in Central Europe, lacking any
conifer species. This pattern repeats itself in Asia from ca. 45° N all the way to the southernmost
occurrence of Pinus merkusii; few species occur in the lowlands and, as in Europe, these are pines
(Pinus). Conifer forests in the mountains of Asia are almost everywhere dominated by species
of the Pinaceae. In North America the montane to subalpine conifer forests of the USA, Mexico
and parts of Central America are likewise dominated by species of Pinaceae. In the E of the con-
tinent a different pattern of distribution prevails from ca. 40° N southwards. Whereas species of
Abies, Picea and Tsuga are confined to the Appalachian Mountains, several species of Pinus are
widespread on the Atlantic Plain where they can form extensive pine-dominated forests or pine
savannas, especially in the S parts of the plain in South Carolina, Georgia, Florida, and along the
Gulf of Mexico. This pattern is continued in parts of the West Indies and Central America. The
Pinaceae arose in the Middle or Late Jurassic (Leslie et al., 2012) when Laurasia and Gondwana
were separated by the Thetis Ocean. However, this lineage seems to have diverged in the Early
Triassic ancestry, and is probably most closely related to the Voltziaceae, now extinct (Farjon,
2008: 71). Major diversification occurred in the Early Cretaceous, but the radiation of most genera
towards the formation of extant species is of much younger age and occurred within the last 25
My (Leslie et al., 2012). We are of course poorly informed about species diversity during earlier
periods, despite a relatively abundant fossil record. As the present distribution of Pinaceae indi-
cates, much of this diversity would have occurred in mountainous regions, and mountains do not
create fossils, they destroy them by continuous erosion.
20 global and trans-continental distributions

Taxaceae

The family Taxaceae is a small family with 24 species and is mostly confined to the Northern
Hemisphere, with two exceptions in the Malesia/SW Pacific regions.

Map GTC-15. The global distribution of Taxaceae—detail.

The family Taxaceae has a wide distribution in the middle latitudes (ca. 25° to 60° N) of the North-
ern Hemisphere, where its species occur in W North America, E North America, Europe, N Africa,
W Asia, the Himalayas and E Asia as far as Japan and the Kuril Islands. Outliers are present in Mexico
and Central America, in Viet Nam, Sumatera, the Philippines and Sulawesi, and most remarkably, in
New Caledonia. Much of this extensive distribution is due to the genus Taxus, including all outlying
occurrences except New Caledonia (map GTC-23 on p. 31). The only other trans-continental genus is
Torreya, with a presence in North America and in E Asia (map MAJ-118 on p. 247). With the excep-
tion of E Canada, the family Taxaceae is absent in the boreal coniferous forest zone and in Canada
it has only a sporadic presence in these northern conifer forests and becomes more common in its
southern transition zone with deciduous angiosperm forest. In Scandinavia Taxus baccata (the only
species of the family in Europe) is mostly restricted to the Atlantic and Baltic coasts (map EM-42
on p. 169). With the exception of T. canadensis (climate zone 2 according to Eckenwalder, 2009)
no species of Taxaceae are adapted either to extremely cold and long winters or to aridity; they
require more mesic climate conditions, which is a limiting factor to their distribution. This probably
explains their absence in continental mountain systems away from oceanic influence, such as the
Rocky Mountains SE of Idaho in the USA and the Tian Shan/Altai mountains in Central Asia. The
presence of Taxus baccata in the Azores (now probably extinct) and on Madeira in the Atlantic
Ocean, and perhaps the presence of T. wallichiana in Malesia as well, is evidence of long distance
dispersal at least in that genus. Can the presence of Austrotaxus spicata, a sister species to the gen-
era Pseudotaxus + Taxus (S. Mathews, unpublished data) in New Caledonia be explained by long
distance dispersal? Estimates of the divergence time of the separation of Austrotaxus and Taxus
place this event at ca. 100 Ma when New Caledonia had not yet separated from Australia and this
continent was part of Gondwana and was situated in high S latitudes, very far away from Laurasia.
Although all unambiguous fossils of Taxaceae have been found (so far) in the Northern Hemisphere,
it is most likely that the family also occurred elsewhere in the Southern Hemisphere.

Podocarpaceae

The family Podocarpaceae is the second largest family with 175 species and with a Southern
Hemisphere/pan-tropical distribution, extending to temperate regions in the Southern Hemi-
sphere and to a lesser extent in the Northern Hemisphere.
 global and trans-continental distributions 21

Map GTC-16. The global distribution of Podocarpaceae—detail.

The family Podocarpaceae has its greatest area of occupancy (AOO) in the tropics, with major
extensions to temperate regions in the Southern Hemisphere and relatively minor extensions
to temperate latitudes in the Northern Hemisphere. As a consequence of its latitudinal range,
the distribution of this family is fragmented, with three main regions, the Americas, Africa and
Southeast Asia/Pacific, separated by oceans. Island distribution is a striking feature of this family,
only surpassed by the Araucariaceae in terms of proportional distribution (map GTC-17 on p. 22).
South America and Central America have a total of 35 species and fewer than 30 occur on other
continents, leaving at least 110 species distributed on islands, with ca. 100 species exclusively so.
The majority of species in Podocarpaceae occurs on islands in the Caribbean Sea, on Madagascar,
in Malesia, the SW Pacific (including New Zealand) and Tasmania; the greatest taxonomic diver-
sity is found in Malesia. Its species have dispersed far into the SW Pacific to Fiji and even Tonga
(on far right and on far left in the map). The Podocarpaceae is predominantly a family of montane
forest species, which is reflected in patterns of its meta-distribution. In the Americas, a more or
less continuous line of distribution runs from Mexico through Central America along mountains,
joining the Andes of South America; an island string of occurrences joins this line in Venezuela.
In the S it is interrupted by a gap (see map SA-1 and discussion on pages 429–430) and then
continues to Tierra del Fuego. Lesser occurrences are found in the tepuis country of the Guiana
Shield and in the uplands of S Brazil. In Africa species of Podocarpaceae are almost exclusively
Afro-montane, coming down to near sea level only in the Cape Provinces of South Africa. In Asia
and the Pacific the majority of species occurs in mountains, but some occur in lowland ‘kerangas’
on nutrient poor soils. In Australia and New Zealand the family is more or less equally divided
between lowland and upland species. The Podocarpaceae date back to the Triassic (Farjon, 2008:
71; Leslie et al., 2012) and originated in Gondwana. The present occurrences in sections of conti-
nents and islands that were once parts of Laurasia or accreted to its tectonic plates are the result
of relatively recent dispersals and radiations. Several species of Podocarpaceae in South America
are more closely related to species in the SW Pacific and Malesia than to some other species in
the Americas (Biffin et al. in Turner & Cernusak, eds., 2011). This indicates independent lineages
repeatedly spreading and diversifying from a southern Gondwanan origin, perhaps Antarctica. In
Malesia the phylogenetic relationships can be complex and are so far hardly researched in the
context of distribution patterns. With a comprehensive phylogeny hypothesis now being devel-
oped (S. Mathews, pers. comm.), this Atlas provides the biogeographical framework for such
research.

Araucariaceae

The family Araucariaceae is a small family with 37 species and a very disjunct distribution, mostly
in Australasia and Malesia, but also in South America.
22 global and trans-continental distributions

Map GTC-17. The global distribution of Araucariaceae—detail.

The family Araucariaceae has three genera, Agathis, Araucaria and Wollemia. Agathis occurs
in Australasia (including New Zealand) and Malesia, Araucaria in Australasia (including New
Guinea) and South America, and Wollemia is confined to one location in New South Wales,
Australia. It appears therefore that the family occurs in the Southern Hemisphere, with some
extensions across the Equator in Malaysia, Sumatera, Borneo and the Philippines. The great
disjunction occurs in Araucaria, which has two species in South America, two in E Australia,
two in New Guinea, one on Norfolk Island and 13 in New Caledonia (map AU-74 on p. 391).
Is this family of Gondwanan origin? Estimates based on molecular dating techniques place its
origin in the Early Triassic ca. 250 Ma (Leslie et al., 2012); fossil evidence goes back to the Late
Triassic and in the Jurassic Araucaria, apparently the oldest of the extant genera, was present
in both Gondwana and Laurasia (Florin, 1963). The fossil evidence of Agathis is younger and
more or less confined to where it still occurs, although more widespread in Australia (Hill & Bro-
dribb, 1999); that of Wollemia, only recently discovered, is limited to Australia and New Zealand
(+ possibly pollen from Antactica) and at most of Late Cretaceous age. Araucaria fossils from
the Jurassic are remarkably similar to some of the extant species; it is a primitive and conserva-
tive genus which has retreated from an almost cosmopolitan distribution to a few refugia in the
Southern Hemisphere. Only in New Caledonia, under special geological and ecological condi-
tions, could a renewed radiation into several species succeed, but these species, due to isolation
on a small island far from major land masses, remained confined to a very small area. Agathis
adapted to tropical and subtropical lowland and montane evergreen forest habitats and spread
to many islands in the SW Paciific and Malesia, but its distribution in Australia became more and
more confined to the narrow strip of coastal rainforest in Queensland that now remains after an
extended period of aridification that turned most of Australia to desert or fire-prone Acacia and
Eucalyptus woodland (White, 1994b). Reconstructing the area of origin of this ancient family and/
or its three genera using phylogenetic inference and the fossil record remains elusive (Givnish &
Renner, 2004) partly because the only location that now has substantial species diversity is the
isolated island of New Caledonia.

Distribution of Genera

Seven of the 70 genera in conifers have a wide and trans-continental distribution and their maps
are presented here. All but one, Podocarpus, are (virtually) confined to the Northern Hemi-
sphere; these are Abies, Larix, Picea and Pinus (Pinaceae), Juniperus (Cupressaceae) and Taxus
(Taxaceae).
 global and trans-continental distributions 23

Pinus
The genus Pinus is with 113 species the largest genus of conifers. It has the widest distribution of
all genera in the family Pinaceae and is nearly congruent with the entire range of the family.

Map GTC-18. The global distribution of the genus Pinus.

In the northern boreal forest zone, the genus Pinus is less widespread than Picea and Larix; its
westernmost localities in North America are in coastal Alaska at the head of Lynn Canal near
Haines and Skagway (Viereck & Little, 1972). In both Canada and Siberia Picea and Larix extend
further N to the Arctic tree line than Pinus, which reaches the tree line in Scandinavia only
because of the influence of the Gulf Stream. Despite these differences, many more dots could
be placed on the map in this zone (shaded) if herbarium specimen data had been available. The
number of species in North America and Central America is much higher than in Eurasia, which
influences the density of dots on the map, however, the general pattern of distribution should be
reasonably accurate S of the 50th parallel. In the USA western pines are separated from eastern
pines by a broad prairie zone which extends from the Gulf of Mexico in Texas to Saskatchewan
and SE Alberta in Canada. Arid and some semi-arid regions in the American West are marked
as having no pines and in the E of the USA there are similar gaps where angiosperm forest pre-
vails (or agriculture replaces it). In Mexico, the land of Pinus in terms of species diversity as well
as forest dominance, the Chihuahuan Desert, the middle part of Baja California (Sonoran Desert),
the Gulf coast in the NE and the Yucatan Peninsula (tropical angiosperm forest) are the only
areas where Pinus is virtually absent. In Europe S of Scandinavia and Scotland (boreal zone) the
genus Pinus is abundant in S Europe and the Mediterranean and in parts of Central Europe, but
virtually absent (naturally) in the European lowlands from France and Ireland to Lithuania and
Belarus. In many areas on sandy soil Pinus sylvestris has been planted and from these plantations
it became naturalized, but all these occurrences are excluded from the map. Eastwards the genus
Pinus extends to the Caspian Sea, but from there to E Afghanistan there is a wide gap, the same
as seen in map GTC-14 for the entire family. The Himalayan chain stands out as it does for the
family as well, and beyond this mountain range Pinus becomes widespread again, but with fewer
species than in North America. Open areas on the map in E China would once have been lowland
or upland angiosperm forest but are now mostly under intense cultivation. Planted and natural-
ized pines are less extensive there than in Europe. The genus Pinus extends S into the tropics
both in the Americas and in SE Asia and reaches S of the Equator in Sumatera. The distribution
is often disjunct at these more southern latitudes, with pines in E and W Cuba but not in between
and in Sumatera and the Philippines but not in Peninsular Malaysia and Borneo. The genus has
reached oceanic islands such as Guadalupe Island off Mexico in the Pacific, the Canary Islands in
24 global and trans-continental distributions

the Atlantic and the Ryukyu (Nansei-shoto) Islands in the East China Sea. The phylogeny of the
genus Pinus has been extensively researched (for reviews see Price et al. in Richardson, ed., 1998;
Farjon, 2005b) but interpretation of the phylogeography of the entire genus will be very complex
and difficult. When the pines of Mexico are marked on a phylogenetic tree derived from complete
plastid genome sequences, they appear in many different clades of American species. American
clades do separate from Eurasian clades at deeper nodes in the tree, but can be nested within
clades at the subgenus level (Parks et al., 2012). Despite this, some observations may be of interest
here. The species Pinus ponderosa, very widespread in W North America (map NA-21 on p. 47) is
sister to a whole suite of Mexican pines that form a well supported clade and thus appear to have
a common ancestor. Apparently, geological and other circumstances induced speciation in this
ancestor in Mexico to a much greater extent than in W North America. The species of subsec-
tion Strobi (Farjon, 2005b) form a well supported clade and divide into two subclades, one with
American species and one with Eurasian species. Looking at the distribution of these, it appears
that this small group of closely related species (‘white pines’ in forester’s parlance) has succeeded
to spread just as widely across the Northern Hemisphere as the genus itself. The next step in this
analysis, calculating the ages of the nodes of the phylogenetic tree, allows us to estimate the speed
at which this radiation may have occurred. According to Leslie et al. (2012) the split between this
clade and its sister, P. krempfii/subsection Gerardianae occurred sometime in the Oligocene to
Miocene (averaged at 23 Ma). In comparison with the age of the genus, Early Cretaceous or ca. 120
My, this radiation took place rather late in its history.

Forest of Pinus nigra ssp. laricio near Bergerie de Ballone, Corsica, France. Characteristic of many pine forests
is their relative open structure, allowing sunlight to reach much of the ground, and their preference for
warmer, S-facing slopes or drier, rockier soil than most other forest forming trees at the same latitude or
altitude. This has enabled the genus Pinus to spread wider and occur further S than other genera in the
Pinaceae. Photo credit Sylvia Philips.
 global and trans-continental distributions 25

Picea
The genus Picea with 38 species is widely distributed in the N parts of the Northern Hemisphere
and is one of the most important forest forming conifer genera.

Map GTC-19. The global distribution of the genus Picea.

The genus Picea is in most areas the dominant conifer genus in the boreal taiga forest and the
shaded areas in this map could be covered almost throughout in dots if the herbarium collection
data had been available. Picea appears to be absent from the most eastern parts of Siberia but
is widespread across the Bering Strait in Alaska. Only five species occur in these forests, three
in North America and two in Eurasia. To the S of the taiga, spruces move up from the lowlands
into the mountains except along the coasts of the N Pacific and N Atlantic Oceans, where some
species due to high rainfall occur from just above the high tide line to variable altitudes in the
mountains. Picea is a northerly genus adapted to low or extremely low winter temperatures and
growing seasons that can be as short as five or six weeks. As a result the species in this genus
at more southern latitudes either occur on coasts with cold waters and much cloud cover, or at
high altitudes. The southernmost occurrences are in Mexico and Taiwan at 23° 20’ N and 23°
13’ N respectively. In many regions spruce forests are often monotonous dark forests consisting
of one or sometimes two species, effectively excluding other trees, although natural forests are
always more varied than plantations in age classes, structure and the formation of gaps allowing
growth of a new generation as well as other plants. The greatest diversity of Picea is found in the
high mountains of W-central China, followed by Japan. Phylogenetic analysis of spruces appears
to indicate that most species evolved during the Neogene and are closely related even across
continents (Leslie et al., 2012). A long suspected example is the very close relationship between
Picea mariana and P. omorika (they hybridize in cultivation) with the former very widespread in
the North American boreal forest (map NA-37 on p. 55) and the latter a relict from the European
Pleistocene now confined to a small location in the Balkan Peninsula (map EM-18 on p. 152). The
sister species to all other spruces appears to be another relict, P. breweriana (map NA-43 on p. 58)
on the California/Oregon border. Both relict species have been found widely in the fossil record,
albeit not outside the continent they now occur in. Although in need of more detailed analysis,
it appears from these results that both range contractions and range expansions have been com-
monplace and have given rise to speciation across wide areas of the Northern Hemisphere. This
process may still be active in China, where several polymorphic species occur together or in close
proximity.
26 global and trans-continental distributions

Taiga forest with Picea glauca in Quebec, Canada. The forest is characterized by the monotony of a
single species of spruce, uneven ages and/or sizes of individual trees, patches of forest destroyed by fire,
and numerous lakes. Downloaded 25 January 2013 from Wikimedia Commons; photo credit Association
PeupleLoup.

Larix
The genus Larix has 11 species; three of which are widespread in the boreal forest zone, with the
remainder in mountain areas of North America, Europe and Asia.

Map GTC-20. The global distribution of the genus Larix.

In the boreal taiga forest of North America, a single species, Larix laricina occurs throughout from
Alaska to Newfoundland. In Eurasia two species occur in the taiga zone, L. sibirica in the W part
and L. gmelinii in the E part. Larix does not occur in Scandinavia but is represented by L. sibirica
from Arkhangelsk on the White Sea coast to the Yenisei River in Siberia, where a zone of hybrid
swarms (L. ×czekanowskii) separates it from L. gmelinii further E. [Herbarium specimen data are
 global and trans-continental distributions 27

very limited for the Russian part of the boreal forest zone.] In E Siberia Larix tends to dominate
the taiga forest instead of Abies and Picea mainly due to increased fire frequency (Schulze et al.,
2012). An isolated population of L. gmelinii occurs in Kamchatka in a relict taiga forest. In the E
of North America L. laricina extends S into the zone of mixed conifer/angiosperm forest as far S
as West Virginia (map NA-47 on p. 61). The genus Larix, like Picea, is adapted to low winter tem-
peratures and short growing seasons, but unlike Picea and other boreal conifers it is deciduous.
The deciduous habit in angiosperm trees is related to this adaptation, but it seems enigmatic that
of conifers only Larix at these N latitudes is deciduous in winter. South of the boreal zone Larix,
like Picea, retreats into the mountains except in the E of North America, where it is also present
in lowland forest. In the USA two species of Larix occur in mountains of the Pacific Northwest
but not elsewhere in the Rocky Mountains or ranges to the W of these. In Europe, just one species
is limited to the Alps and the Carpathians, but with a few presumably remnant localities in the
Polish lowlands, indicating a more extensive distribution in the past which, as Late Pleistocene
pollen records show, extended on to the plains of Central Europe. In Asia, the genus Larix occurs
much further S than in North America and Europe and extends S to the Longchuan-Salween
Divide in Yunnan at 25° 30’ N. Several species are distributed in this region from the E Himalaya
(Central Nepal) through the high mountains of NW Yunnan, Sichuan and S Gansu, where they
reach altitudes well above 4000 m. Further occurrences are in the mountains of NE China, North
Korea and Japan, and in the Altai Mountains and E Tian Shan of Central Asia. In a conifer-wide
phylogenetic analysis that sampled seven of the 11 species (Leslie et al., 2012) the inferred relation-
ships among the species are not congruent with their continental distribution.

Taiga forest with Larix sibirica in autumn colour, Picea obovata and Betula sp. near Gornoknyazevsk, close
to the River Ob in the Yamal region of W Siberia. Of these three species, Betula grows fastest and Picea
slowest and the dynamics of forest fires determines the composition of the taiga forest. The lower this
frequency, the more Picea obovata will dominate, eventually replacing both birch and larch. Photo credit
and © Bryan Alexander/Arcticphoto.com.
28 global and trans-continental distributions

Abies
The genus Abies has 47 species and is widely distributed in the Northern Hemisphere, from the
northern boreal forest zone to most of the major mountain systems in more southern latitudes.

Map GTC-21. The global distribution of the genus Abies.

Only two species of Abies are part of the boreal taiga forest, A. balsamea in North America and
A. sibirica in Asia; none occur in the boreal zone in Alaska or in Scandinavia. The shading of the
taiga forest zone on the map indicates where they occur in Canada and Russia, but neither spe-
cies extends as far N as Larix and Picea. To the S of this zone, Abies is widely distributed in most of
the major mountain systems N of the Tropic of Cancer and extends S to El Salvador and Honduras
in the Americas and to N Viet Nam and Taiwan in Asia. In North America S of the taiga zone the
distribution of Abies is widely separated in E and W sections and it is the W section that extends
through Mexico to tropical Central America as far S as 14° 21’ N. Gaps in this W section occur and
are related to regions, including mountain ranges, with insufficient precipitation, particularly in
the ‘Basin and Range Province’ between the Rocky Mountains and the Cascades/Sierra Nevada.
Deserts limit Abies in the SW of the USA and in N Mexico. In the E the Appalachian Mountains
are the southernmost refugium, as indeed here Abies fraseri occurs on isolated summits in relict
populations. In Europe one species is widespread from the Pyrenees to the Balkan Peninsula;
other species are relicts, with some of the rarest in the genus on mountains in the Mediterra-
nean. Turkey, Lebanon and the Caucasus Mountains are areas with species of Abies in W Asia. E
of these occurrences there is again the gap from the Caspian Sea to E Afghanistan also observed
for Picea, Pinus and Taxus, where mountains and deserts are too dry for these conifers. Abies
has a limited distribution in Central Asia in the westernmost Tian Shan and to the E in the Altai
Mountains. Its major occurrence starts in the Hindu Kush and extends through the Himalayas
far into China, with greatest abundance and diversity in the high mountains of NW Yunnan, W
Sichuan and S Gansu. Another important region for this genus is in NE China, Korea and Japan.
Isolated populations (and species) are present in N Viet Nam, SE China and Taiwan and some of
these represent the rarest species of the genus. The phylogeny of Abies has been studied by sev-
eral researchers (e.g. Isoda et al., 2000b; Xiang et al., 2004; 2009; Leslie et al., 2012), with a common
result that A. bracteata, a morphologically aberrant species from California, is sister to the rest of
the genus. Some emerging clades are congruent with regions of distribution, e.g. the species of
Europe, Mediterranean, Turkey and Caucasus, but most are a mix of Asian and North American
species, perhaps to be interpreted as the result of several migrations and subsequent radiations.
A complex of species certainly occurs in the Sino-Himalayan region, where morphological poly-
morphism has given rise to divergent species concepts among taxonomists. Similar problems
occur with species of Abies in Mexico (Jaramillo-Correa et al., 2008), where evidence from DNA
is incongruent with that from morphology and the biogeography seems complex.
 global and trans-continental distributions 29

Forest of Abies amabilis and a few trees of Picea engelmannii in Wenatchee National Forest, North Cascades,
Washington, USA These trees form narrow spires typical for environments in which snow cover is both
heavy and of long duration, to minimize breakage of branches. Photo credit Aljos Farjon.

Juniperus
The genus Juniperus with 53 species has the widest distribution of all conifer genera but is limited
to the Northern Hemisphere except in Africa, where one species crosses the Equator.

Map GTC-22. The global distribution of the genus Juniperus.

In the boreal forest zone a few decumbent species of Juniperus are widespread, in particular
J. communis var. saxatilis which occurs across Eurasia and in NW North America (maps EM-31-
32 on pages 160–161) and J. horizontalis in North America (map NA-80 on p. 81). The former is
the only native conifer in Greenland and Iceland. S of the boreal forest zone junipers are as
widespread as the map indicates due to two properties of this genus, a wide range of habitat
adaptations but particularly aridity, and dispersal of the seeds by birds. The latter property has
taken junipers to far off islands such as the Azores, Bermuda and the Ogasawara Group of the
Bonin Islands. Junipers can be prostrate shrubs in subarctic or alpine tundra or boreal conifer
forest, or tall trees in forests that receive abundant rainfall. Many species survive in the margins
30 global and trans-continental distributions

of deserts and in steppes where rainfall is sporadic or highly seasonal. In regions where deciduous
or semi-evergreen angiosperm forest is the dominant vegetation (now mostly converted to agri-
culture) the genus Juniperus is naturally sparse, limited to locations with eroded topsoil such as
inland sand dunes and heaths. These places are often anthropogenic, the result of deforestation
and overgrazing many centuries ago. The spread of Juniperus communis in lowland NW Europe
is an example of this, but it may have happened in prehistoric times several millennia ago. We
count these as natural occurrences (see photograph on p. 162). In some regions, e.g. the SW USA
and W-central Asia, junipers are dominant or co-dominant trees or large shrubs in open, semi-arid
woodlands which can cover very large areas. In other parts of the world, they can be large trees
occurring in mixed conifer forests, e.g. in the Sierra Nevada of California, in the Taurus Mountains
of Turkey, in the highlands of Ethiopia and Kenya, and in SE Xizang [Tibet] and E Qinjiang in
China. In some regions, these forests were cut long ago, as in N-central China, but many juniper
species are pioneer trees invading secondary, open woodland and abandoned grazing lands. In
the USA, where this is well documented, the spread of J. occidentalis in the W and J. virginiana in
the E are examples of range expansions in recent historic time with anthropogenic causes. Other
species do not have this capacity and are therefore more sensitive to logging; J. procera in Africa
belongs to these. It is the only species with a distribution that extends S of the Equator, but it
may be on the retreat path, with the southernmost occurrence a single mature tree (map AF-5 on
p. 462). The genus Juniperus divides into two subgenera, with the smallest made up of species that
occur mostly in Eurasia (including the Mediterranean) with the exception of the circumpolar spe-
cies J. communis. The largest subgenus is nearly as widespread as the genus, but within it there are
some clades with continental distributions, e.g. North America (incl. Mexico) and Asia (Adams,
2011; Leslie et al., 2012). Among the latter is a clade that contains the scale-leaved junipers with
almost black ripe seed cones and a single seed per cone. These species groups radiated during
the Neogene, presumably from an ancestral species that occurred in the same continent. Other
phylogenetic relationships among species are more complex and may have involved migration,
subsequent radiation, and even hybridization.

Open woodland/steppe in Arizona, USA with Juniperus deppeana (tree) and J. monosperma (shrubs). Some
trees of the former species (known as Alligator juniper for their bark pattern) are very large and presumably
old, indicating ancient woodland converted to grassland but now being invaded by the much younger
shrubs of J. monosperma. Photo credit David Thomburg.
 global and trans-continental distributions 31

Taxus
The genus Taxus has 10 species with a wide distribution across the temperate regions of the North-
ern Hemisphere, but with some outlying populations in the tropics N and S of the Equator.

Map GTC-23. The global distribution of the genus Taxus.

The genus Taxus is distributed in several well separated regions in the temperate latitudes of
the Northern Hemisphere and in some isolated locations in the tropics, in Malesia on either side
of the Equator. In North America yews occur in three regions, the Pacific Coast of the USA and
Canada from California to Alaska and the NW Rocky Mountains between latitudes 46° 35’ N and
51° 30’ N, E North America with an outlying population in N Florida, and Mexico, with outly-
ing populations in Guatemala and Honduras. The fourth region is Europe, Morocco and Alge-
ria, Turkey and the Caucasus, with outlying populations in the Azores, Madeira and the Elburz
Mountains in N Iran. The fifth region extends from the Hindu Kush in E Afghanistan through the
Himalayas to NE India, Myanmar [Burma], China, N Viet Nam and Taiwan. The sixth region is
in NE China, Primoriye and Sakhalin Island in the Russian Far East, Korea and Japan, extending
to the southernmost Kuril Islands. One species, which has its westernmost population in Central
Nepal, occurs well beyond its main range in isolated populations in S Viet Nam, the Philippines,
Sumatera and Sulawesi and crosses the Equator in the two latter islands. The major disjunctions
in the global distribution of Taxus are related to climate. Yews are trees or shrubs growing under
mesic conditions of temperature and humidity, often but not exclusively under canopy of other
trees (they can themselves shade out all vegetation). Most species do not withstand long periods
of extreme cold and are therefore absent from the boreal forest zone as well as from regions with
a continental climate; the exception appears to be T. canadensis with occurrences in the boreal
forest of E Canada. In the Himalayas both the relatively low latitude and the monsoon influ-
ence from the E, which allows Taxus to occur deep into Sichuan and beyond, have a mitigating
influence on the severity of the winters. That yews occur in the tropics seems remarkable; the
altitude range there is wide, 1200–3000 m, but not extreme and within the range of many tropi-
cal conifers. The phylogeny of Taxus (Li et al., 2001; Leslie et al., 2012) is probably still in need of
further work, but some interesting results from these analyses can be mentioned. The first is that
Taxus baccata and T. contorta (syn. T. fuana in the latter analysis) are sister species in a clade
with two other sister species, T. canadensis and T. cuspidata. The entire fourth region (Europe
etc.) mentioned above has one species, T. baccata (map EM-42 on p. 169); T. contorta occurs in
the W Himalayas and Hindu Kush (map MAJ-32 on p. 194). Given the nature of the gap in terms
of climate and its width, as well as the circumstance that routes of migration of seed eating
birds do not cross it but pass in a N↔S direction, this may represent vicariance (a single species
range separated to become two species). Taxus canadensis is the widespread species in E North
32 global and trans-continental distributions

America, T. cuspidata the species of the sixth region (Japan and environs). This biogeographical
pattern is well known from many genera of trees and is best explained as another example of
vicariance. Taxus chinensis and T. wallichiana (incl. T. sumatrana) also form a clade and here the
nature of the distribution, with scattered outliers on islands related to a large and more contiguous
distribution in mainland Asia, would suggest that dispersal events have played a significant role
in the distribution and separation of these species.

Looking down Kingley Vale towards the ancient yew wood, Sussex, England. Taxus baccata grows in pure
stands on chalk with very thin soil, or is mixed with angiosperms, often Fagus sylvatica (beech) on the
upper slopes and in the vale where soils are deeper. Photo credit and © James Giles/Wild Images.

Podocarpus
The genus Podocarpus is the second largest genus of conifers with 97 species. This genus has
a pan-tropical/Southern Hemisphere distribution with extensions N of the Equator into warm
temperate regions in Mexico, China and Japan.

Map GTC-24. The global distribution of the genus Podocarpus.

 global and trans-continental distributions 33

The distribution of the genus Podocarpus is similar to that of the family Podocarpaceae (map
GTC-16 on p. 21). The main differences are the absence of Podocarpus in Ethiopia (where it is
replaced by Afrocarpus, see map AF-11 on p. 465), and in S India, where only Nageia wallichiana
occurs (map MA-55 on p. 323). In Chile Lepidothamnus fonkii is the southernmost member of the
family, but almost everywhere else where other genera of the family occur, Podocarpus is also
found. Not only is Podocarpus the most widespread genus by a wide margin, in many regions it is
the only genus, enlarging the extent of occurrence (EOO) of the family as a whole; examples are
Mexico, the West Indies, SE and S Brazil, Madagascar, E Himalaya, Taiwan, Japan, Tonga and SW
Australia. The distribution of the genus, due to its latitudinal extent lying in the tropics and the
Southern Hemisphere, is naturally divided into three regions, the Americas, Africa/Madagascar
and Asia/Australasia. In terms of geographical extent the Americas and Asia/Australasia are
about equal and Africa/Madagascar is much smaller; in terms of species diversity Asia/Australasia
exceeds the other two combined. With a few exceptions, e.g. in SW Australia and in coastal flats
of Borneo, the distribution of Podocarpus follows mountain systems, e.g. from Mexico through
Central America and along the Andes in South America. Where the genus occurs on islands these
are usually also mountainous. Tropical forested lowlands are mostly areas without any species of
Podocarpus, examples are the Yucatán Peninsula in Mexico, S Nicaragua, parts of Borneo and New
Guinea, and most importantly, the large basins of the Amazon and Congo Rivers. Phylogenetic
analyses of the genus are still progressing in terms of efforts to increase the scope of taxa and gene
sequences sampled, as this genus to date has been studied much less intensively than e.g. Pinus.
Some interesting results already emerge (Leslie et al., 2012). One clade contains all species of Tas-
mania, New Zealand and SE Australia plus one, P. gnidioides, from New Caledonia. The other New
Caledonian species appear in a well separated clade, which is sister to a clade with NE Australian
species plus two species from Malesia which are very widely distributed across that region. This
would imply two different instances of radiation, from two different ancestors possibly in New
Caledonia, divergence between which is associated with the first split among extant lineages and
is dated back to the Palaeogene, when New Caledonia was much closer to the Australian con-
tinent than at present. The four African species in the genus form a clade, indicating common
ancestry, but no species from Madagascar were included in this wide ranging sample of conifers.
The relationship of the five Malagasy species with the four existing on the continent will be of
much interest. As is the case with other large conifer genera, diversification of the genus to its
present level seems to have occurred in the Neogene, i.e. during the last 25 My, from a common
ancestor that arose in the Palaeogene.

Tropical montane evergreen forest with Podocarpus milanjianus as an emergent tree on Mt. Bigugu,
Nyungwe National Park, Rwanda, Central Africa. The species of Podocarpus are co-dominants at most and
do not form mono-specific stands but are distributed as scattered individual trees in a diverse forest. Photo
credit and © Coke Smith (www.cokesmithphototravel.com).
NORTH AMERICA

Geography

The continent of North America in the Atlas of the World’s Conifers comprises Canada and the
continental United States of America, i.e. Alaska and the ‘lower 48’ states, plus Greenland. Of
Greenland, only some small ice-free coastal areas in the southern part are relevant to conifers and
in the statistics that follow Greenland is not included; no conifer taxa occur there that do not also
occur in North America. North America (N of Mexico) has a total land area of 19.27 million km².
A large part of this territory is situated in the High Arctic beyond 70° N where no conifers occur;
this includes Baffin Island and the Canadian Arctic Islands as well as the Arctic continental coast-
line from the Beaufort Sea in Alaska to Hudson Strait between Baffin Island and the N coast of
Quebec. The most striking topographical division of North America is the mountainous West,
extending from Alaska into Mexico, and the lowland central and eastern parts of the continent,
only interrupted in the E by the lower and much more ancient Appalachian Mountains. Hud-
son Bay and the large lakes in Canada and the USA, all the result of expansive Pleistocene ice
caps, form large bodies of water in the N and NE of the continent. Nearly the whole of Canada
is dotted with numerous lakes across a broad boreal forest zone stretching across the continent
from the Atlantic to the Pacific. In contrast, and this is the second major division, the SW of the
continent is not only mountainous but also dry, with extensive areas classified as deserts. Nearly
flat lowlands surround the Gulf of Mexico and extend northwards along the Atlantic coast on the
E-side of the Appalachian Mountains. The interior of the continent, from southern Saskatchewan
to Texas, was an expanse of treeless plains (before agriculture) dissected by rivers that drain
into the Missouri-Mississippi River system, the largest drainage in the continent. Eastwards, the
prairies gradually merged with deciduous angiosperm-dominated forests. Westwards they still
rise into the foothills of the Rocky Mountains, with the ca. 60 million bison completely replaced
(but reintroduced) by European cattle. The Rocky Mountains are a broad and complex system
of geologically young mountain ranges rising to 4401 m (Mt. Elbert) with many other summits
in Colorado and Wyoming over 4000 m. The highest mountains are situated in Alaska and the
Yukon Territory, with Denali (Mt. McKinley) at 6194 m the highest mountain in the continent.
Between the Rocky Mountains in the USA and the Pacific Ocean lie a series of N-S oriented
mountain ranges and interspersed basins, culminating in the Sierra Nevada of California with
Mt. Whitney at 4421 m. In Canada and Alaska, the Rocky Mountains lie adjacent to the Coast
Ranges, the latter in part submerged in the Pacific Ocean forming an extensive archipelago. Cli-
matically, the continent is diverse, ranging from extreme arctic in N Greenland and the Canadian
Arctic to subtropical in Florida and from the cold and wet coasts of Labrador and Newfoundland
to one of the hottest and driest spots on earth, Death Valley in California, dropping to 86 m
below sea level. Extensive cool temperate rainforests dominated by conifers occur in the Pacific
Northwest of the USA (W Oregon, W Washington), coastal British Columbia and the Alexander
Archipelago in S Alaska.
 north america 35

Conifers in North America

Families Genera Species Species + infra Countries with Area of occupancy Endemic taxa
conifers in km²
3/8 17/70 99/615 126/794 2/2 106,850/19,269,000 113
Numbers after ‘/’ are global or continent totals.

Few species occur in the boreal forests of Alaska and Canada, but they are widespread and abun-
dant. Herbarium collection data are inadequate to cover the occurrence of these conifer species
and as a result there are far too few dots on the map; the white areas between them would be
filled too if the specimens had been available. Collections of herbarium specimens are particu-
larly few in Quebec and Ontario around Hudson Bay (Larix laricina, Picea glauca, P. mariana)
and in N Saskatchewan (the same species + Pinus banksiana). The density in California reflects
the high number of species there, not a continuous conifer forest, and the same applies to other
areas with many dots on the map. The accuracy of the map is much greater in the contiguous
United States, i.e. S of the Canadian border. There are several striking patterns to be observed.
The richest areas for conifers are in California around the empty Central Valley, and N into the
western halves of Oregon and Washington. Some areas of the Rocky Mountains stand out, in
Idaho-W Montana, Wyoming and Colorado into N New Mexico. In the eastern part of the con-
tinent the southern Appalachians, Florida and the Mexican Gulf Coast to the east, and an area
from Virginia to New Hampshire stand out as rich in conifer species. The Midwestern states are
more thinly populated and conifers are very scarce in the (former) prairie states, with large parts
of these states devoid of conifers. North Dakota, South Dakota, Nebraska and Kansas; also large
parts of E Montana, Minnesota, Iowa and Missouri, as well as S Texas, have few if any conifers. In
the western USA, Nevada has large areas without conifers, as do substantial parts of the Mojave
Desert in California and the Sonora desert in Arizona. In the eastern USA, the original dominance
of deciduous angiosperm forest (now experiencing a come-back with the decline of agriculture
especially in New England) limited the extent of conifers, which probably explains the many
blank areas. Aridity is the main constraint further west, where tree growth in general, not just
conifers, is limited to banks of rivers, or in deserts virtually absent. At high altitudes conifers are
more abundant although trees such as aspen (Populus tremuloides) often grow alongside them
covering large areas. In the far west the montane forests are nearly pure conifer forests, often with
a relatively high number of species. Fire-prone habitats that allow tree growth are also mostly
dominated by conifers and are common in California but also in the boreal forests of the N and
in certain parts of the Rocky Mountains.

Map NA-1. Distribution of conifer species in North America (N of Mexico) and Greenland.

36 north america

Conifer diversity in North America (N of Mexico) is substantial at the level of genera and species,
but only represented by three families, Cupressaceae, Pinaceae and Taxaceae. The 17 genera are
divided among the families as follows: nine in Cupressaceae, six in Pinaceae and two in Taxaceae.
The genera in Cupressaceae are Calocedrus, Chamaecyparis, Cupressus, Juniperus, Sequoia, Sequoia­
dendron, Taxodium, Thuja and Xanthocyparis. In Pinaceae the genera are Abies, Larix, Picea, Pinus,
Pseudotsuga and Tsuga. In Taxaceae they are Taxus and Torreya. Of all these, only two are endemic
to North America, Sequoia and Sequoiadendron, each with a single extant species. Like most of the
other 15 genera, these two were widespread in the Northern Hemisphere before the ice ages of the
Pleistocene. There is indeed a strong element of circum-boreal distribution in the North American
conifer flora, represented by genera such as Abies, Larix, Picea, Pinus, Juniperus and Taxus. Other
genera represent an East Asian/North American distribution pattern that is well publicized for
angiosperm trees since the days of Asa Gray in the 19th century. These are Calocedrus, Chamaecy­
paris, Thuja, Xanthocyparis, Pseudotsuga, Tsuga and Torreya. The genus Cupressus is present in W
North America (including Mexico), Europe (Mediterranean, North Africa to Iran) and East Asia to
the Himalayas. Apart from the two sequoias, only Taxodium is no longer found in Eurasia. It occurs
in the eastern USA and in Mexico and Guatemala. Endemism is much higher at species level and
when infraspecific taxa are included, only 13 taxa from a total of 126 occur across the border with
Mexico and none are shared with the West Indian islands. Nine of these cross the border into Baja
California from a greater extent of occurrence (EOO) in California, USA. While the actual bor-
der between Mexico and the USA is obviously an artificial line, in view of the substantial conifer
diversity in both countries (see the chapter Mexico, Central America and the West Indies) there is
obviously a somewhat diffuse but yet marked biogeographical border in terms of conifer species
diversity. In Baja California Norte, nearly all species are northern, just extensions of their ranges. In
mainland Mexico, the species of the USA are replaced by those of Mexico and Central America. In
terms of species diversity, the two most diverse genera in North America are Pinus with 38 species
and Juniperus with 14 species. Whereas most genera occur in both the eastern and western halves of
the continent, at species level and S of the boreal zone there is a marked division. In the contiguous
USA no species are distributed right across the continent, although some species are closely related
and are perhaps vicariants, i.e. a population that became separated into an eastern and a western
part which then evolved separately into different species. Examples are Pinus strobus (E) and
P. monticola (W) and Juniperus virginiana (E) and J. scopulorum (W). The division between E and
W is well demonstrated in map NA-2 below, where the central prairie states, where not devoid of
conifers, contain only Cupressaceae (= Juniperus) and Pinaceae and Taxaceae are widely separated
in an eastern and a western section. In the contiguous USA both east and west have their own dis-
tinct species. This virtually complete taxonomic separation at species rank suggests a lengthy geo-
graphical separation. The American Great Plains were an inland sea until the end of the Cretaceous.
Tropical lowland forests dominated during the Eocene and grassland steppes from the Miocene
onward. Only a few junipers occasionally occur in this vast region now, most of them in Texas.

Map NA-2. The distribution of species in Pinaceae (black), Taxaceae (yellow) and Cupressaceae (red) in
the contiguous United States.
 north america 37

In North America as defined here, the state of California is a centre of diversity, with 15 out of 17
genera and 52 species or nearly 53% of the total number for the continent in an area only 2.1%
of the landmass. Diverse genera in California are Pinus with 18 species and Abies and Cupressus
with seven species each. Of the genus Cupressus this is 47% of the world total of 15 species (Far-
jon, 2010a). Endemism in California is also high, with Sequoiadendron giganteum confined to the
state and Sequoia sempervirens nearly so; in addition there are the following endemic species:
Abies bracteata, Cupressus goveniana, C. macnabiana, C. macrocarpa, C. sargentii, Pinus torreyana,
Pseudotsuga macrocarpa and Torreya californica. With just one tiny stand of Pinus balfouriana
on the Oregon side of the state border, this species is nearly endemic to California. To the N the
state of Oregon is second in diversity, with 14 genera and 32 species. There is substantial overlap
between the two states and Oregon has no endemic species. Of course the boundary between
these two states is arbitrary too, and a few species e.g. Chamaecyparis lawsoniana and Picea brew­
eriana have limited distributions that straddle the border. In fact these occur in a small area
centered on the Klamath Mountains with 14 genera and 34 species, the most diverse limited area
for conifers in North America. Diversity as can be expected diminishes further N while endemism
as exemplified by the last two species disappears completely N of the 43rd parallel. The E side
of North America has markedly fewer species, with the highest numbers in Florida, six genera
and 13 species. Taxus floridana is endemic to the state, but Torreya taxifolia, although similarly
limited in its distribution, occurs also in the neighbouring state of Georgia. As on the west coast,
diversity diminishes further north such that in the whole of Canada + Alaska there are no more
than 11 genera and 35 species, many of which only just cross the (again artificial) border with the
contiguous USA. The continental boreal conifer forest, which covers the greatest portion of the
landmass of Alaska and Canada, contains just 11 species of conifer in five genera.
Although California and the Klamath Mountains clearly stand out as the (partly overlapping)
centres of diversity for conifers in North America, their levels of endemism (ca. 20% for Califor-
nia) are too low to treat them as separate sections in this Atlas. The species in this chapter are
mapped and discussed in taxonomic order, but with distribution patterns as a secondary criterion
for grouping them. The sequence of families and genera follows the one presented in Christen-
husz et al. (2011) to reflect phylogenetic relationships.

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Map NA-3. Map of North America showing numbers of conifer genera per one degree cell.

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Map NA-4. Map of North America showing numbers of conifer species + infraspecific taxa per one degree cell.
38 north america

Maps NA-3 and NA-4 show that the areas with the greatest taxonomic diversity of conifers are
California and the Pacific Northwest of the USA. A lesser centre of diversity, particularly at genus
level, is situated in the New England states and adjacent parts of Ontario, Canada. In the West,
the area with high generic diversity is larger than the area with high species diversity. The latter
is more concentrated in California and especially N California + SW Oregon and this is also the
region with the top number of 11–12 genera per degree cell. These are the Klamath and Siskiyou
Mountains on either side of the California-Oregon state line, the most diverse region for conifers
not only in North America but in the Western Hemisphere.

Map NA-5. Rarity values for 1 degree cells. Cell score is the total of the rarity calculations for each species
in the cell. The rarity calculation is 1 divided by the total of cells occupied by a species. Thus, a species that
occurs in one cell scores 1 while a species that occurs in 10 cells scores only 0.1.

Map NA-5 shows local endemism to be mostly concentrated in California and SW Oregon. Two
species, Taxus floridana and Torreya taxifolia are responsible for the red 1 degree cell in NW
Florida. The Pacific Northwest, while scoring high in diversity, is low on local endemism and the
same is true for the New England area. Conifer taxa in these regions all have wider distributions.
North America is thus exceptional among the continents and other super-regions considered in
this Atlas in having its local endemism of conifers concentrated in one area. This area largely
coincides with the California Floristic Province, characterized by endemism in many other plant
groups. A combination of mountain ranges forming connections to especially more northern
regions, isolation of other mountains especially in S and coastal California, a varied topography
and climate and the presence of the fog-bringing cold California Current have caused this ende-
mism to develop (see Debreczy & Rácz, 2011 vol. 2: 550–563 for a detailed overview).

Pinaceae

The Pinaceae is the most prominent family of conifers in North America, both in terms of num-
bers of species and in terms of forests, as they make up the majority of forest forming conifer
trees. A few species of Abies, Larix, Picea and Pinus make up the vast boreal conifer forests in
the N of the continent and the conifer forests in the Rocky Mountains. The same genera, with
different species, dominate along the Pacific coast, in the Cascade Range and the Sierra Nevada,
but here, further south, other conifer taxa in Cupressaceae and Taxaceae are also present. In the
E of the USA pine forests become prominent in the lowlands of North Carolina, South Carolina,
Georgia, Florida, Alabama and Missisippi.

Pinus

There are 38 species of Pinus in North America; these amount to 33.6% of all species in the genus.
The distribution of the genus in North America is similar to that of the family as a whole (see
 north america 39

map NA-2 on p. 36 for the contiguous USA) and is not shown here. The species are grouped here
showing similarity in distribution patterns rather than taxonomic affinity, with a few exceptions
where appropriate.

Map NA-6. Pinus banksiana [n=115, m=113, h=15, 1747–2010]

Pinus banksia is the only species in the genus with a nearly cross-continental distribution. It
occurs in the boreal forests from Nova Scotia and E Quebec to the Mackenzie River, but further
W it is absent in British Columbia, S Alaska and the Yukon Territory, where it is replaced by Pinus
con torta (map NA-20 on p. 47). Its southernmost outlier is in Indiana and another one slightly
further N is in New Jersey. These are effectively outposts into the deciduous broad-leaved forest of
the E USA; its usual habitat is dry sandy areas, sometimes fringes of peat bogs, in the cold boreal
forest. Pinus banksiana is well adapted to fires, which occur frequently in the short but warm and
dry summers, quickly seeding out from its serotinous cones that do not fall from the branches
and open after the heating by fire. In early successional stages it often dominates, but later other
species of Pinaceae move in, e.g. Larix laricina, Picea glauca and P. mariana. These conifers partly
out-compete the pines, until the next fire destroys them. Pinus banksiana reaches the arctic tree
line in the N of Canada, but remains well away from Hudson Bay and is also absent in N Quebec.
For these and other northern regions the herbarium specimens found are too few and far apart
to produce an adequate map of the true distribution. Critchfield & Little (1966) published a range
map of the species that better shows the extent of occurrence.

Map NA-7. Pinus resinosa [n=61,

m=61, h=5, 1876–2010]

Pinus resinosa is widespread in

the region of the Great Lakes,
SE Canada and New England.
It occurs from the SE corner of
Manitoba and N Minnesota east-
ward to Nova Scotia and (rarely)
in Newfoundland. It has a few
southern outliers in Illinois, Indi-
ana and Virginia, but its main distribution lies in the boreal forests within the latitudes where
pure conifer forests merge with deciduous angiosperms such as Acer, Betula, Fagus and Quercus.
In the north of its range conifers dominate, in the south angiosperms. This area was heavily
40 north america

glaciated during the last Ice Age and the glaciers left huge deposits of sand, some of which in post-
glacial time was blown into sand dunes. It is to these sands that this pine is mostly confined, as
here it can successfully compete with other trees. It is largely a lowland pine growing at altitudes
between 200 m and 450 m, but ascends to 1300 m a.s.l. in the Appalachian Mountains. This pine
was heavily exploited in colonial times and into the twentieth century, it is now making a come-
back in forests that are secondary growth and often managed in a more restorative way.

Pinus strobus var. strobus near Jefferson, Maine.

This is secondary forest gradually turning back to
the type of mixed angiosperm-conifer forest which
covered much of the state before logging began.
Photo credit Aljos Farjon.
Map NA-8. Pinus strobus var. strobus (black) [n=100,
m=99, h=10, 1880–2009]; P. strobus var. chiapensis
(red) n=70, m=68, h=26, 1900–1994]

Pinus strobus has an extremely disjunct distribution. By far the largest range of the species is in
E North America, but other populations occur far to the S in Mexico and Guatemala. Pinus strobus
var. strobus occurs from the SE corner of Manitoba and SW Ontario through Minnesota, Wiscon-
sin and NE Iowa, then E as far as Newfoundland; along the Appalachian Mountains it reaches
S to N Georgia and Tennessee. Isolated populations exist in Illinois, Indiana and Kentucky. In the
N part of its eastern limit it follows the Atlantic coast, but S of Chesapeake Bay it retreats inland,
leaving the Atlantic Plain for the Appalachian Mountains. Pinus strobus var. chiapensis occurs
in small and often isolated populations in Puebla, Veracruz, Guerrero, Oaxaca and Chiapas of
Mexico and in El Quiche and Huehuetenango of Guatemala. This seems a strange disjunction,
but among angiosperms there are several species of trees with a similar distribution. The key to
this disjunction is the migration route for northern trees along the Appalachian Mountains and
around the Gulf of Mexico that functioned during glacial maxima in the Pleistocene, enabling
these species to escape into Mexico. The climate in the lowlands around the Gulf is now subtropi-
cal and unsuitable, as it was in earlier inter-glacial periods. But it was similar to the climate in
present-day Massachusetts and New Jersey during the ice ages, where Pinus strobus occurs in the
lowlands up to the coast. The southern populations were ‘left behind’ in the relatively cool and
certainly wet mountain climate of S Mexico and Guatemala. The altitude range of var. chiapensis
is 500–2200 m and no frost is known to occur in any part of its range. This variety is not hardy,
so certainly some genetic sifting has occurred (natural selection) to produce trees well adapted
to this different environment.
 north america 41

Map NA-9. Pinus monticola [n=108, m=104, h=13, 1825–2009]

Pinus monticola is closely related to Pinus strobus and

is is possibly the western element of a vicariance event
that split the ancestral species in two, now separated
by ca. 1500 km between the westernmost occurrence
of P. strobus (map NA-8 on p. 40) and the eastermost
localities of P. monticola. The distribution of P. mon­
ticola divides into two ranges. The most extensive
of these is from Vancouver Island and the opposite
coast of British Columbia along the Cascade Range
southward as far as Tulare Co., California in the Sierra
Nevada. The eastern range is in the Rocky Mountains
of SE British Columbia, NE Washington, N Idaho and
just across the border with Montana. Very scattered
occurrences are reported from NE Oregon (Critchfield
& Little, 1966) but herbarium collections are not avail-
able. In Oregon P. monticola is also rare in coastal areas, only nearing the ocean more abun-
dantly where the Klamath Mountains extend westward. In California it retreats to the high Sierra
Nevada. Limits of precipitation and perhaps high summer temperatures determine this pattern;
in the Sierras it can be found above 3000 m but in interior British Columbia not above 500 m.
Pinus monticola can colonize disturbed areas and for a time form pure stands, but eventually
many other conifers, depending on location and site conditions, will join it to form a diverse
mixed conifer forest. This species is susceptible to Western white pine blister rust (Cronartium
ribicola, Basidiomycota); ‘old growth’ conifer forests with mixed species in all age classes are far
less susceptible to mass outbreaks than the young, even-aged, species-poor stands that prevail
under active forest management for exploitation of timber.

Map NA-10. Pinus lambertiana [n=103, m=94, h=30, 1825–2010]

Pinus lambertiana is related to P. monticola as well as

to P. strobiformis and perhaps other species in sub-
section Strobi; the phylogenetic relationship of taxa
within this group is still not well resolved (Leslie et al.,
2012; Parks et al., 2012). Pinus lambertiana has a range
that partly overlaps with P. monticola but it has a more
southerly distribution; it extends from Clackamas Co.
in Oregon, USA to Baja California Norte in Mexico.
It is most abundant and attains its greatest size (70–
75 m tall, 3–3.8 m d.b.h.) in the Cascade Range and Sis-
kyou Mountains of S Oregon and in the Sierra Nevada
of California. In the Sierras it is mostly confined to
the western slopes, but extends E of the divide into
Nevada around Lake Tahoe. South of this mountain
range this pine becomes more scattered and limited
to the highest mountains in S California and finally
the Sierra San Pedro Martír in Baja California. A few
small, isolated stands of P. lambertiana occur in the
Santa Lucia Mountains along the Pacific coast S of Monterrey, California. In the Cascade-Sierra
Nevada Range P. lambertiana is a major component of the famous mixed conifer forest which is
one of the best developed of its kind in the world, with many species attaining great size, among
them the Giant Sequoia of California.
42 north america

Map NA-11. Pinus jeffreyi [n=119, m=115, h=26, 1850–2009]

The distribution of Pinus jeffreyi is very similar to

that of P. lambertiana, especially in California. In
Oregon it is confined to the Klamath Mountains in
the SW corner of the state. In Baja California Norte,
Mexico it occurs in the Sierra de Juarez as well as
in the Sierra San Pedro Martír, where this species
reaches its southern limit. The main range of P. jef­
freyi is in the Sierra Nevada and the high mountains
of S California into Baja California. There are a few
isolated stands in the Coast Ranges, more in the N of
California than in the S of the state, where the sum-
mers become increasingly dry and hot. Pinus jeffreyi
is another major component of the mixed conifer for-
est that dominates the middle elevations of the Sierra
Nevada W slopes. This pine extends in many places
across the divide and is also present in the Lake Tahoe Basin across the border with Nevada. Few
herbarium collections from Nevada seem to exist, so the map is incomplete there.

Map NA-12. Pinus edulis [n=95, m=84, h=11, 1846–2008] Map NA-13. Pinus quadrifolia [n=72,
m=69, h=20, 1850–2008]

Pinus monophylla in the dry hills fringing the Mojave Desert in

Map NA-14. Pinus monophylla [n=139, southern California. Photo credit Aljos Farjon.
m=134, h=28, 1843–2008]
 north america 43

The three species of maps NA-12-14 are commonly known as pinyon pines, from the Spanish
piñon or nut, referring to the edible seeds these species produce. Two have distributions that
extend across the border into Mexico, but either their main range or a substantial portion of it
is in the USA. Other species and subspecies of pinyon pines occur (mainly) in Mexico and are
mapped and discussed in the chapter Mexico, Central America and the West Indies. Pinus edulis
has an extensive range largely in the ‘Four Corner’ states of Utah, Colorado, Arizona and New
Mexico. A significant population occurs in the New York Mountains at the E margin of the Mojave
Desert in California and there are a few scattered localities with this species in S Wyoming and in
‘Trans-Pecos’ Texas. In Arizona the Mogollon Rim is its SW limit. Pinus edulis is the pinyon pine
of high altitude plains, mesas and slopes in the semi-arid region of the Colorado Plateau and its
environs. It forms extensive, open stands commonly with one or more species of Juniperus known
as Pinyon-Juniper woodland, which is one of the most widespread semi-arid vegetation types
in North America. Pinus monophylla occurs W of P. edulis in W Utah, Nevada and California; its
northernmost outlying populations are in S Idaho, while southward it extends into Baja Califor-
nia. The two species are sympatric in parts of Arizona, but P. monophylla is less common there
(a taxon P. edulis var. fallax described from Arizona is here included in P. monophylla). While
P. monophylla is most widespread in Nevada, where it forms open pinyon-juniper woodland on
every mountain range (it is the Nevada State Tree) there are few dots on the map in the central
part of that state because apart from NY few herbaria consulted have specimens. Pinus quadrifolia
has a limited distribution in S California, USA and N Baja California, Mexico. Its northernmost
locality is in the Coachella Valley, Riverside Co. and its southernmost locality is near the Socorro
placer mines above Rancho San José in the foothills of the Sierra San Pedro Martír, a total N-S
distance of ca. 400 km. This species occupies a zone between desert and lower altitude chaparral
and the high altitude mixed conifer forest (of the highest mountains) in pinyon-juniper woodland
but also in more shrubby vegetation, with this the only small tree. All pinyon pines are adapted
to semi-arid conditions with hot summers but at high altitudes deep snow cover often occurs in
winter, especially in more northern latitudes.

Map NA-15. Pinus balfouriana (black)

[n=39, m=39, h=9, 1852–1992]; P. longaeva
(red) [n=54, m=54, h=14, 1898–2005];
P. aristata (yellow) [n=56, m=53, h=5,
1861–2008]

Pinus aristata on Windy Ridge in the Rocky Mountains Pinus balfouriana in Sequoia National Park, Sierra
SW of Denver, Colorado. Photo credit Mena Schmid. Nevada, California. Photo credit Aljos Farjon.
44 north america

The three species of map NA-15 are the ‘foxtail pines’ so called because the 5-needle fascicles
are long retained on the branches, giving the appearance of bushy tails. They are more famous
for their longevity, with the record age for a single individual tree (not a clone) held by a live
specimen of Pinus longaeva which was >4900 years old when it was felled on Wheeler Peak,
Nevada in 1964 (Lanner, 1999). In the White Mountains of California 17 trees were found to be
>4000 years old, the oldest tree there having 4600 annual rings. This tree, named Methuselah
and growing in the Schulman Grove named for its discoverer, was still alive more than 50 years
after this count but its locality and identity are now concealed for fear of souvenir hunters with
axes and saws. The other two species can attain a few millennia too, but not as many years as P.
longaeva. These pines are very closely related and botanically similar, yet their distributions do
not overlap. As a group (clade) they are sister to the Pinyon pines discussed above. Pinus longaeva
has a wide distribution from E California (E of Owens Valley) through Nevada and Utah, where
it usually occupies summit areas (to 3700 m) of mostly N-S oriented mountain ranges and the
highest escarpments and mesas of the Colorado Plateau and the granite batholith mountains that
protrude from it such as the Henry Mountains. It often grows on bare scree slopes and in rock
outcrops, alone or together with P. flexilis. Pinus balfouriana is known from two disjunct popula-
tions (which have been given taxonomic distinction as varieties or subspecies) in California. The
northern population is scattered in the Klamath Mountains and consists of mainly small, isolated
stands on S-facing slopes between 1600 m and 2400 m. The southern population occurs in the
Sierra Nevada, mainly within Kings Canyon and Sequoia National Parks but with a few stands
in Inyo and Tulare Counties E and S of these famous national parks. It occurs at higher altitude,
from 2700 m to 3700 m. Pinus balfouriana can grow on various rock types, in the Sierras mainly
on granite, often bare of other vegetation apart from some patches of ground-covering shrubs.
It grows ‘pure’ or mixed with a few other conifers, e.g. Pinus con torta, P. albicaulis and Juniperus
occidentalis. Pinus aristata is the easternmost species in this group, mainly occurring in the Rocky
Mountains of Colorado and N New Mexico at 2500–3400 m altitude. An isolated population is
known from the San Francisco Mountains N of Flagstaff, Arizona. There is some concern that
climate change may negatively affect recruitment (through increased competition from other
conifers). However, it has to be realised that during the millennia through which some of these
trees have lived the climate has changed several times, while the uneven age of individuals in the
populations is evidence that they successfully recruited all that time.

Pinus flexilis among limestone rocks on Sheep

Mountain, Rocky Mountains, Colorado. These gnarled
trees have established in a rock fall that happened
centuries ago and are able to survive on the poorest
of substrates where rainfall and snowmelt will drain
away almost instantly. Photo credit Mena Schmid.

Map NA-16. Pinus flexilis [n=125, m=118, h=17,

1861–2008]
 north america 45

The two varieties recognized under P. flexilis, var. flexilis and var. reflexa (which latter takes the
species into Mexico) are here not distinguished on the map. Pinus flexilis is widely distributed
in the Rocky Mountains and in the Sierra Nevada and other mountains in S California. It is also
present in the mountains of the Great Basin in Nevada and Utah, on the high sections of the
Colorado Plateau, and in Arizona (var. reflexa) where it may ‘grade’ into P. strobiformis (map
MCW-26 on p. 106). US foresters and botanists treated the latter species without consideration
of Mexican populations and much of what they mapped as that species [Critchfield & Little,
1966 map 8, repeated in Flora of North America 2: 381 (1993)] is better considered to belong to
P. flexilis (Farjon & Styles, 1997). Pinus flexilis is a high mountain species which is found as high
as 3800 m in New Mexico, but comes down to 900 m in the Black Hills of South Dakota. It is also
present in other isolated and elevated locations E of the Rocky Mountains in Montana, North
Dakota and Nebraska. This species is one of several in the genus Pinus of western North America
that can withstand extreme conditions of climate on bare rock or scree without any other veg-
etation cover. On these sites it occurs either alone or with Pinus albicaulis and Abies lasiocarpa
in the northern parts of its range and with Pinus aristata in the SE and P. longaeva in the SW. At
lower altitudes it is usually only a minor component of more diverse conifer forest. Pinus flexilis
is highly susceptible to infection and death from the disease white pine blister rust caused by the
non-native pathogen Cronartium ribicola. Infections have been known since the 1960s and appear
to be spreading from N to S causing large scale death of trees in many areas.

Map NA-17. Pinus albicaulis [n=123, m=122, h=27,

1862–2009]

Pinus albicaulis is a high mountain pine

with a more northerly distribution than that
of P. flexilis, but in large areas their ranges
overlap and they can occur together, some-
times causing identification problems in the
field if no seed cones are available. Isolated
stands have been observed at ca. 54° N in
British Columbia and the main range starts
at ca. 53° 20’ N; the southernmost occurrence
is in Tulare Co., California at 36° 33’ N. In
Alberta and British Columbia Pinus albicau­
lis occurs in the mountains to ca. 2300 m; all
collections at higher points were made in the
USA, up to 3700 m above Bullfrog Lake in the
southern Sierra Nevada, California. In British Columbia (Canada) and Washington (USA) there is
a wide gap between populations in the coastal mountains and in the Rocky Mountains. Although
widespread in the Rocky Mountains, this species has only been collected at one locality in Colo-
rado, Estes Park (J. S. Newberry s.n., US61140, the holotype of the species). This is now recognized
as the easternmost outlier population of P. albicaulis. Apparently in the high Rocky Mountains
of Colorado and New Mexico P. albicaulis is replaced by P. flexilis (map NA-16 on p. 44). In the
W it follows the Cascades-Sierra Nevada Range, with greatest abundance in the highest parts of
the Sierras. In the ‘intermountain region’ between the Rocky Mountains and the Cascades-Sierra
Nevada Range isolated populations are present on the highest summits and ridges in Oregon and
Nevada. More than any other pine in North America, P. albicaulis is characteristic of tree line
alpine habitat, where it often forms dense thickets of windswept dwarf trees.
46 north america

Map NA-18. Pinus torreyana subsp. torreyana (black)

[n=5, m=5, h=3, 1875–1982]; P. torreyana subsp. insularis
(red) [n=3, m=3, h=3, 1900–1992]

Pinus torreyana is an extremely rare species of

which three small populations are known, one on
the mainland California coast N of San Diego near
Del Mar and two on the Channel Islands of Santa
Catarina and Santa Rosa. The island populations
are recognized as the distinct subspecies insularis.
Near Del Mar, most trees are within Torrey Pines
State Park, but a smaller subpopulation occurs to the N and scattered trees are outside the protected
area, where urbanization is encroaching on their habitat. An estimated total of 3,500 trees of P.
torreyana subsp. torreyana still exist. On the two islands together there are about 1,000 trees. This
species is dependent on the cooling effect and moisture (fog) brought in from the ocean during sum-
mer almost every day. Its habitat, sparse chaparral and occasionally a few oak trees in ravines on the
islands, is easily burnt especially with the increased risk from urban development and tourism to the
islands. Both subspecies are listed in the category Endangered (EN) on the IUCN Red List.

Map NA-19. Pinus radiata var. radiata (black) [n=21, m=21,

h=11, 1827–2007]; P. radiata var. binata (red) [n=40, m=40,
h=13, 1875–1987]

Pinus radiata is another relict species on the California

coast and on two islands off Baja California, Mexico.
The mainland population, known as P. radiata var.
radiata, is separated into three small subpopulations,
the largest of which is on the Monterey Peninsula and
in Point Lobos State Park and a few points S along
the coast. The other two are to the N in Santa Cruz/
San Mateo counties (mostly in Santa Cruz County)
and to the S around Cambria in San Luis Obispo Co.
Pinus radiata var. binata occurs on Guadalupe Island,
240 km offshore, and on Cedros Island, nearer the
coast of Mexico. Pinus radiata is in its natural habitat confined to promon tories and strips of
rocky coast as well as two islands; it is rarely found more than 11 km from the sea. Here it thrives
within the fog belt (which also daily engulfs the islands), associated with Cupressus macrocarpa
and Pinus attenuata in Monterey and with Quercus tomentella on Guadalupe Island. It is the
nominate variety from the mainland that has turned out to be such a vigorous plantation tree in
New Zealand, South Africa, Chile and elsewhere in the Southern Hemisphere that now millions of
hectares are covered with this pine, dwarfing its natural population. In California itself, where it
has been planted as an amenity tree rather than a production tree, it has naturalized throughout
the western parts of the state, making a mockery of its original biogeography. The obvious ques-
tion arises: why is its natural distribution so confined if it is so successful (it is an invasive species
in most countries where it has been introduced for plantation forestry). The full answer will be
complex and is not yet fully understood. Poor seedling establishment through competition from
chaparral shrubs as well as frequent fire outside the fog belt play a role in natural ecosystems,
but when people started bulldozing for roads and urban development, these limiting factors were
diminished. Once carried outside its confines by planting, the species could spread. This became
even more dramatic when Radiata pine was taken to continents where competitors were either
absent or had been removed (deforestation) and no pathogenic checks existed. Almost ironically,
Pinus radiata in its natural habitat is listed as Endangered (EN) on the IUCN Red List, with the
mainland variety most at risk of extinction.
 north america 47

Map NA-20. Pinus con torta var. con torta (red)

[n=56, m=49, h=13, 1794–2008]; P. con torta var.
latifolia (black) [n=124, m=122, h=12, 1869–2008];
P. con torta var. murrayana (yellow) [n=49,
m=47, h=22, 1825–2007].

Pinus con torta is widely distributed in the

Rocky Mountains and in the Coast Ranges
of Alaska and British Columbia, extending
southwards along the coast of Washington,
Oregon and N California as well as through
the Cascades/Sierra Nevada Ranges. Three
varieties are recognized with largely allo-
patric (distinct) ranges. Pinus con torta var.
con torta is distributed along the Pacific
coast from Alaska to California, including
the Alexander Archipelago, Queen Charlotte Islands and Vancouver Island. Pinus con torta var.
latifolia occurs from the Yukon Territory through the Rocky Mountains S to Colorado and N Utah;
also further west in British Columbia, Washington and Oregon. Outlying (disjunct) populations
are known from the Black Hills in South Dakota, Cypress Hills Provincial Park, Saskatchewan
and the Liard Mountains in the Northwest Territories; these are all shown on the map. There are
very few herbarium collections from the northern parts of its range in Canada and our map is
incomplete for this region. Pinus con torta var. murrayana has the smallest of the three ranges and
occurs in the Cascades Range of S Oregon and N California, the Sierra Nevada of California, the
Transverse Ranges in S California and in the mountains of Baja California, Mexico as far S as the
Sierra San Pedro Martír. In the northern parts of its range and in parts of the US Rocky Mountains,
e.g. Yellowstone National Park, Pinus con torta can form extensive pure forest, largely because for-
est fires prevent slower growing species of conifer such as Picea and Abies attaining dominance.
In the western mountain ranges this species is more often a non-dominant constituent of mixed
conifer forests, but here, too it reacts to fire by invading the burnt forest areas. Along the coast,
P. con torta is most commonly present on sites with little soil or on podzolic sands or peat. The
altitude range of the species is great, from near sea level along the coast to 3400 m in the Sierra
Nevada and southern Rocky Mountains.

Map NA-21. Pinus ponderosa var.

ponderosa (black) [n=93, m=93,
h=22, 1826–2010]; P. ponderosa var.
scopulorum (red) [n=130, m=128,
h=16, 1846–2008]

Pinus ponderosa is one of the

most widespread pines in North
America and a dominant species
in the pine forests and mixed
conifer forests of the American
West. Its main distribution is in
the USA, but it extends NE into
British Columbia and S just across
the border with Mexico in a few
places. The general distribution of
this species forms a great and nearly closed circle where it is abundant to ubiquitous, with large
areas in the centre in Nevada, SE Oregon, S Idaho, SW Montana, N Utah and W Wyoming where
Pinus ponderosa is absent or very scarce. In some parts this is caused by lack of precipitation,
48 north america

e.g. in Nevada and SE Oregon/SW Idaho, but this is not the cause in W Wyoming (e.g. Yellowstone
National Park and mountains to the S and E) and SW Montana. Critchfield & Little (1966) called
this absence “inexplicable” but there may be combinations of lack of summer rainfall at lower
altitudes and continental cold (short growing season) at higher altitudes at play that favour ‘har-
dier’ species such as P. con torta (map NA-20 on p. 47) and P. flexilis (map NA-16 on p. 44) in this
region. Ponderosa pine is usually divided in a Pacific variety var. ponderosa and a Rocky Mountain
variety var. scopulorum. These two varieties are well separated by the central gap but merge in the
northern part of their ranges where it is not always easy to separate them taxonomically. The mild
and wet coast of Washington and British Columbia, including Vancouver Island, is unsuitable for
this light and sun requiring pine, and only a few isolated and small populations are known, e.g.
on the N coast of the Olympic Peninsula (see map) and in the rain shadow of the mountains W of
Puget Sound. Even the W-slopes of the Cascades in Washington and Oregon are generally too wet
for Ponderosa pine. This variety is at its most imposing in the Sierra Nevada of California, where
trees can reach 70+ m tall and 2.5 m d.b.h. The very widespread Rocky Mountain var. scopulorum
is most abundant in the mountains and scarce on the Colorado Plateau in the ‘Four Corners’ area
where the states Arizona, Utah, Colorado and New Mexico meet. Critchfield & Little (1966, map
47) painted extensive ‘blobs’ there indicating large forests or stands, but the relevant herbaria, e.g.
RM and ASU, have no collections from this region, so it is presumably rare there, too. It probably
occurs in some of the batholiths e.g. the Henry Mountains and Abajo Mountain. Probably the
furthest E on to the Great Plains is a small stand near Bassett, Nebraska (red circle). In Mexico,
an interesting outlier of P. ponderosa var. scopulorum occurs on a mountain near the Cañon de
Sentenela, Sierra del Carmen, Coahuila. Pinus ponderosa (var. arizonica) in Mexico on map 47 in
Critchfield & Little (1966) is Pinus arizonica and its varieties (map MCW-21 on p. 103), a species
that crosses the US-Mexico border.

Pinus ponderosa var. ponderosa in dry hills on the

Columbia River, Washington, USA. This is probably
not far from where David Douglas collected seeds
and specimens in 1826 that formed the basis of its
original description by C. Lawson in 1836. There
is no unambiguous type specimen; a small shoot
collected by Douglas at Kettle Falls on the Columbia
is kept at K but there is no evidence that Lawson
saw it. Despite this, it is very likely that these trees
represent Douglas’s species and that any concept of
P. ponderosa in a narrow sense (i.e. var. ponderosa)
has to include them. This means that if the much
taller trees in Oregon and California are thought to
be different from these taxonomically, they cannot
be called Pinus ponderosa var. ponderosa. Photo
credit Aljos Farjon.
 north america 49

Map NA-22. Pinus attenuata [n=73, m=69, h=23, 1852–2007]

Pinus attenuata is endemic to the California Floris-

tic Province (CFP), which includes 70% of California
and parts of SW Oregon and N Baja California. This
region has a Mediterranean type climate with hot, dry
summers and mild, rainy winters. The CFP is rich in
plant species, many of which are endemics, includ-
ing several conifers. Pinus attenuata occurs in a belt
around California’s Central Valley and is more scat-
tered in N California and S Oregon, where it reaches
its northernmost point in Lane Co. In S California it
is found in the San Bernardino Mts. and San Jacinto
Mts. as well as near the coast and in Baja California
only near Ensenada on the coast. Its altitude range is
250–1700 m, so it occurs in a colline to low montane
open forest zone. This species is well adapted to for-
est fires, which are frequent in this habitat. Extremely
tenacious and serotinous seed cones only open after
heating by fire to release the seeds. This pine forms pure stands or is mixed with cypresses and
oaks, or occurs in chaparral, especially in more southern sections of its range. Fires give it a
chance to regenerate, without these periodic removals of shrubs and trees it could not compete
with other trees that would then prevail.

Map NA-23. Pinus muricata [n=54, m=52, h=20, 1832–1997]

Pinus muricata is a strictly coastal species distributed

intermittently along the coast of California from Hum-
boldt Co. in the N to Santa Barbara Co. in the S; also
on the Channel Islands Santa Rosa, Santa Cruz and
Santa Catalina. In Baja California, Mexico there are a
few localities N of Ensenada, mainly W of San Vicente
and near Eréndira and San Isidro. The pines on Cedros
Island, off the coast of Baja California, are not P. muri­
cata (Critchfield & Little, 1966 map 59) but P. radiata
var. binata (map NA-19 on p. 46). Along this entire
length of Pacific coast, fog rises off the ocean around
late morning to midday in summer and envelops the
vegetation onshore. This has a cooling and humidi-
fying effect allowing these pines to grow here. Just a
few km inland the cooling effect of the cold California
Current which causes the fog to develop is lost and the hot sun burns down on the earth. Pinus
muricata still hangs on to its coastal habitat because of the California Current. Fossil evidence of
its cones in Pleistocene deposits has demonstrated that in cooler times it occured much further
inland (Axelrod, 1983, 1986). Coastal development and related hazards are causing the decline of
many subpopulations and this species is listed as Vulnerable (VU) on the IUCN Red List.
50 north america

Map NA-24. Pinus sabiniana [n=46, m=45, h=11, 1826–2009]

Pinus sabiniana is another endemic species of the

California Floristic Province. Formerly thought to be
endemic to California (Critchfield & Little, 1966; Grif-
fin & Critchfield, 1972) it is now also known from two
localities in Jackson Co., Oregon (one specimen at
ORE was already collected in 1945 but only correctly
identified by Aaron Liston of Oregon State University
in 1999; the other collection at OSC is from 2006).
This species occupies a similar range and habitat as
P. attenuata (map NA-22 on p. 49) in a broad belt
around the Central Valley from 50 m to 1800 m a.s.l.
It forms extensive open pine woodland in the Coast
Mountains and in a foothill to lower montane zone
on the W flank of the Sierra Nevada. Another outlying
population was found in 2007 along Pine Valley Creek
in San Diego Co. and this is the southernmost locality
now known (specimens at MO and RSA). A curious gap appears in Tulare Co. along the foothils
of the Sierras, already present in map 54 by Critchfield & Little (1966) compiled from mainly
observational data (USDA Forest Service) and not filled by herbarium specimen data accessed
for this Atlas.

Map NA-25. Pinus coulteri [n=62, m=60, h=21, 1832–2008]

Pinus coulteri is famous for its massive seed cones

armed with hooked umbos resembling raptor claws.
It is another endemic of the California Floristic Prov-
ince, with a more southerly and westerly distribution
than P. attenuata and P. sabiniana. It occurs from the
San Francisco Bay area S to the Cañon San Ramón
in the Sierra San Pedro Martír, Baja California. It is
confined to the Coast Ranges and does not occur in
the Sierra Nevada. Its altitude ranges from 300 m to
2150 m, with the higher occurrences in the S parts of
its range. Pinus coulteri is a constituent of southern
Californian mixed conifer or conifer-oak woodland,
merging with chaparral at lower altitudes and with
denser conifer forest of the Canadian Vegetation Zone
at its highest limits. Two pines with which it often occurs (both more abundant than P. coulteri
in most localities) are P. sabiniana (map NA-24 on p. 50) at lower altitudes and P. jeffreyi (map
NA-11 on p. 42) at higher altitudes.

Map NA-26. Pinus taeda [n=96, m=95, h=9,

1832–2010]

Pinus taeda is one of the lowland pines of the

SE United States with an extensive distribu-
tion from the Cape May Peninsula in New
Jersey to Bastrop State Park in Texas. An out-
lying northern population has been found in
W Kentucky. Pine forests are absent in much of
the Mississippi Valley so there is a gap between
 north america 51

the western part of the range and the more extensive eastern range. In Florida Pinus taeda occurs
only in the N half of the peninsula. While it is most common on low sandy ridges and old river
terraces in the Atlantic Coastal Plain, it ascends into the foothills of the Appalachian Mountains
to ca. 700 m. Often forming pure stands as a result of spreading after disturbances of earlier veg-
etation, or more recently on to abandoned farmland, this species is also present in mixed pine
forests or with angiosperm trees in later phases of forest succession.

Map NA-27. Pinus echinata [n=97, m=95, h=10,

1832–2006]

Pinus echinata is widespread in the SE

United States from the Atlantic Coastal
Plain to the foothills and valleys of the
Appalachian Mountains and westward to
the hills of E Oklahoma and the Ozark Pla-
teau of SE Missouri and S Illinois. Along
the coast it extends from New Jersey to E
Texas. A gap occurs in the Valley of the
Mississippi River separating a smaller
western range from a larger eastern range, as with P. taeda (map NA-26 on p. 50). The maximum
altitude at which this pine grows is ca. 600 m and it is uncommon below 150 m, so there is a strip
along much of the Atlantic and Gulf coasts, including nearly all of Florida, where P. echinata is
absent. The map presented here is incomplete mainly due to limited numbers of herbarium col-
lections from the Atlantic Coastal Plain and from E Kentucky, West Virginia and Virginia. Pinus
echinata is often associated with P. taeda but occurs generally in somewhat drier habitat.

Map NA-28. Pinus virginiana [n=88, m=88, h=9, 1847–2009]. Pinus virginiana in secondary vegetation in the
foothills of the Appalachian Mountains in Virginia. Photo credit Aljos Farjon.

The distribution of Pinus virginiana is situated midway between southern species such as P. glabra
and P. palustris and northern species like P. strobus var. strobus. In the S it is separated from the
closely related species P. clausa by ca 150 km (in herbaria, some specimens from Florida identi-
fied as P. virginiana belong to P. clausa, see map NA-32 on p. 53). Pinus virginiana occurs near the
coast in the pine barrens of New Jersey but further southward it retreats from the Atlantic Coastal
Plain to the Piedmont. It is widespread through the Appalachian Mountains and on to the Cum-
berland Plateau and elevated regions of N Alabama and adjacent parts of Georgia. It is a lowland
pine that is usually found below 650 m but has been collected at 950 m on Table Rock in South
Carolina. In many parts of its natural range P. virginiana appears to be a vigorous pioneer tree
invading abandoned farmland. In more stabilized habitat it retreats to poor soils (pine barrens)
52 north america

and rocky terrain with thin soils. If left to its natural course, the pine invasion of fields will prob-
ably succeed to mixed forest dominated by angiosperms where this pine will again only play a
minor role in the vegetation.

Map NA-29. Pinus rigida [n=83, m=83, h=7, 1830–2010]. Pinus rigida in Acadia National Park, Maine, USA.
Photo credit Aljos Farjon.

Pinus rigida occurs in the NE of the United States from coastal Maine through the Appalachian
Mountains to NE Georgia. In the N of its range it is common in sandy areas on the coast and on
islands, e.g. Cape Cod, Long Island and the pine barrens of New Jersey. It also occurs inland, the
farthest N is just inside Canada in Quebec Province at 45° 20’ N. In the Appalachian Mountains its
distribution is intermittent; although herbarium records accessed for the Atlas are not complete,
it is clear that it is rare in Virginia, West Virginia, Kentucky and Tennessee and more common in
W North Carolina and adjacent sections of South Carolina and NE Georgia. Pinus rigida is usually
restricted to infertile soils, in the N of its range glacial outwash plains and sandy shores and islands,
in the S mainly in shallow soils on mountain ridges, steep slopes and sandstone plateaux. The alti-
tude ranges from near sea level to 1370 m, in the N of its range usually below 500–600 m.

Map NA-30. Pinus palustris [n=68, m=66, h=9,

1832–2010]

Pinus palustris is a lowland species occurring

on the Atlantic Coastal Plain of the SE United
States, ranging from SE Virginia to Florida
and westward to E Texas. In the Piedmont it
ascends to about 700 m. On the plains it may
form extensive pine forests, often mixed with
P. echinata and P. taeda, in the foothills of the
mountains it is more often an occasional tree
on thin soil, mixed with Quercus spp. This species has a ‘grass stage’ as a seedling; this is an adap-
tation enabling it to survive ground fires. In the first years of establishment there is little or no
elongation of the stem and only primary leaves develop; the seedling develops an extensive root
system instead. When above ground parts are burnt, leaves quickly appear again. After several
years of building the root system, an erect stem develops without branching, raising the foliage
above the next fire. Branching can then take place without much risk. The strategy works if fires
are frequent and light, as they will be under natural circumstances. Preventing fires, as foresters
are inclined to do, causes a fuel load to be accumulated which then burns too hot and kills the
sapling pines. Prevention of fires also increases competition from herbs and shrubs as well as other
pines, as the seedlings of P. palustris do not initially grow in height. Pinus palustris was once the
most common pine in the Atlantic Coastal Plain, perhaps covering 25 million ha. Exploitation for
 north america 53

timber and naval stores and conversion to farmland and pasture as well as favouring other pines
reduced this to less than 1.6 million ha in 1985. Foresters and landowners prefer other species that
do not delay initial growth with a grass stage and the decline is ongoing, although slowing down.
Pinus palustris is listed as Endangered (EN) on the IUCN Red List.

Map NA-31. Pinus elliottii var. elliottii (black)

[n=43, m=43, h=8, 1869–2007]; P. elliottii var.
densa (red) [n=22, m=20, h=8, 1901–1997]

This is the southernmost species of Pinus in

North America (as defined here) in which
two varieties are recognized. Pinus elliottii
is closely related to P. caribaea, the near-
est population of which occurs on Grand
Bahama Island (map MCW-71 on p. 133);
indeed especially the southern variety of
P. elliottii was in the past confused with P.
caribaea. Pinus elliottii var. elliottii occurs on the Atlantic Coastal Plain from southern South Caro-
lina to SE Louisiana, including N Florida. Pinus elliottii var. densa occurs along the two coasts and
in the centre of peninsular Florida to its southern end, including eight of the Florida Keys. Pinus
elliottii can form pure stands, or mixed pine forest with P. taeda and P. serotina. The var. densa
has a seedling ‘grass stage’ while the var. elliottii lacks it.

Map NA-32. Pinus clausa [n=41, m=40, h=9,

1882–2006]

Pinus clausa is one of the southernmost

species of Pinus in the SE United States,
largely confined to Florida but with a few
localities in S Alabama. This species of
pine occupies the drier areas away from
swamps and eutrophic rivers. It is a fire-
successional species, in particular on infer-
tile white sand dunes and flats of marine
origin and at low elevations between 5 and
60 m a.s.l. or up to 90 m on the west coast. Its vernacular name, Sand pine, indicates the soil on
which it predominantly grows, both inland and along the coasts.

Map NA-33. Pinus glabra [n=47, m=45, h=9,

1872–2008]

Pinus glabra occurs on the southern Atlan-

tic Coastal Plain from South Carolina to
Louisiana. In Florida it is mainly present
in the north of the state, but isolated popu-
lations have been found in coastal locali-
ties further south. Unlike P. clausa this
species is often found along streams and
in hummocks and swamps on wet sandy
soil, not in dry places. It establishes itself
in the shade of broad-leaf trees such as Magnolia, Liriodendron, Liquidambar, Nyssa, Carya, Fagus
and Quercus and survives by overtopping them. This tolerance to shade is rather unusual for a
54 north america

pine and it is diminished as it grows taller, so it often requires some canopy opening to attain
dominance. Unlike many other pines it is susceptible to fire. It also occurs in pine forests with P.
elliottii and P. taeda but in these forests it is only a minor constituent. Where the forest is regularly
flooded Pinus glabra gives way to Taxodium distichum.

Map NA-34. Pinus serotina [n=39, m=38, h=8, 1854–2010]

Pinus serotina is another pine species of the Atlantic

Coastal Plain (seven species of Pinus occur there in
total), reaching the Piedmont in Georgia. This spe-
cies is distributed from New Jersey to Florida and
Alabama. Its habitat is coastal swamps and wet
alluvial flats in the interior where the water table
remains high. In so-called pocosins, peat swamps
that have risen above the water table and drain
outwards, P. serotina is common (a local vernacu-
lar name is Pocosin pine). Pinus serotina is often
associated with Taxodium distichum or with other
pines, e.g. P. taeda and P. elliottii.

Map NA-35. Pinus pungens [n=45, m=45, h=9, 1876–2007]

Pinus pungens is a species distributed in the Appa-

lachian Mountains, from Pennsylvania to South
Carolina and the NE corner of Georgia (Hickory
Nut Mountain). An outlying population is reported
from E Kentucky (Critchfield & Little, 1966) but is
not mapped here for lack of a herbarium speci-
men. Although most occurrences are from the
main ranges of the Appalachians, especially the
Blue Ridge Front in the S part of its range, numer-
ous outlying populations (some mapped here) occur on the E side and some on the W side of this
mountain system proper. These are often on so-called monadnocks, formations that rise higher
than the surrounding Piedmont. The altitude range of P. pungens is from ca. 300 m to its highest
location in the Great Smoky Mountains at 1760 m near the southern end of its range. This pine
usually occupies ridges and steep rocky slopes with thin soils over acidic rock, where the angio-
sperm forest is low and thin, thus escaping competition.

Picea

The genus Picea has seven species in North America; two of these occur across the continent in
the north, the other four have more limited (but in some cases still wide) distributions. This is
a genus of colder climates and as a result it is absent in the lowlands at lower latitudes; in addi-
tion no species have been able to adapt to low levels of precipitation. Its ecological amplitude is
consequently limited and this is reflected in its distribution in North America and to some extent
explains the low species diversity as well.
 north america 55

Map NA-36. The distribution of the genus Picea in North America (N of Mexico).

The boreal forest of Alaska and Canada, where three of six species occur, are poorly represented
on the map due to lack of sufficient herbarium collections for this vast area. South of 49° N (the
western half of the Canada/USA border) the map becomes more accurate. Here Picea is confined
to a narrow strip of the Pacific coast as far S as N California and inland to high mountains in the
W and to the much cooler maritime climate of the NE, until there too it retreats into the Appala-
chian Mountains and finally at its southern extremity to the summits of the highest parts. In the
far W even very high mountains such as the Sierra Nevada of California, with many summits over
4000 m, can be devoid of spruces, although this particular instance may be more of a ‘historical
accident’ than an ecologically determined phenomenon. Picea does extend through the Rocky
Mountains into Mexico but becomes very rare and limited to a few isolated localities (see the
chapter Mexico, Central America and the West Indies).

Map NA-37. Picea mariana [n=166, m=166, h=15, 1872–2009]

Picea mariana is a boreal species distributed across the continent from W Alaska to Newfound-
land. It extends to the Arctic tree line which runs from the southern foothills and valleys of the
Brooks Range in Alaska through the Mackenzie Delta, the northen shores of Great Bear Lake to
the W shore of Hudson Bay, then around that bay to Ungava Bay in N Quebec. Its southern limits
are near Calgary in Alberta, Canada in the W, and in Wisconsin and Connecticut in the USA in the
E. Both the Arctic tree line and the southernmost occurence of Picea mariana are considerably
further S on the E side of the continent than on the W side. In the W this species reaches the
coast furthest S on the Kenai Peninsula, Alaska at ca. 60° N, in the E it occurs furthest S near the
coast in Connecticut at 41° 22’ N. In winter, sea ice comes S along the Atlantic coast as far as Nova
56 north america

Scotia at ca. 45° N while the Pacific coast of Alaska remains ice free. Picea mariana occurs mostly
in bogs or swamps and on permafrost sites (‘muskeg’), at elevations between <150 m and 800 m,
occasionally in western mountains to 1500 m or 1800 m a.s.l., on a variety of acid soils, often on
peat, in the south predominantly so. Except for some of the southernmost stands, P. mariana is
a small, spindly tree, often standing not quite upright due to lateral movement of the permafrost
top layer in summer when it thaws. Although extensively sampled here, herbarium specimens
cannot represent adequately the ubiquitous occurrence of this species over its vast range, with
actual numbers of trees in the trillions. In particular, the Canadian provinces of Saskatchewan,
Manitoba, Ontario and Quebec are poorly represented and could have many more dots if the
remote interiors of these provinces had been better surveyed botanically.

Map NA-38. Picea glauca [n=142, m=142, h=19, 1849–2009]

Picea glauca along Beaver Creek, Yukon

Territory, Canada. Photo credit Aljos
Farjon.

Picea glauca has a very similar gen-

eral distribution to that of P. mariana
(map NA-37 on p. 55), but in the west-
ern USA it is present in a few localities
in Montana and in the Black Hills of
South Dakota. Like P. mariana it is a
boreal forest species extending from
W Alaska to Newfoundland across
the continent, marking the Arctic
tree line in the N and bordering on the northern deciduous angiosperm forests in the SE of its
range. Also like P. mariana, it stays away from much of the Pacific coast but borders the Atlantic
coast from Labrador to Maryland. Picea glauca is a major constituent of the boreal conifer forest,
occurring on better drained sites than P. mariana but not shunning permafrost soils. It is mainly
a lowland conifer but in the W of its range it can be found into the E-side of the Rocky Mountains
of Alberta to ca. 2000 m. Here it merges with the distribution of P. engelmannii (map NA-39 on
p. 57) and probably hybridizes; P. glauca var. albertiana (here included with the species) is pos-
sibly the product of hybridization. As with P. mariana, herbarium collection data are inadequate
to cover the vast range of this ubiquitous species and northern parts of Saskatchewan, Ontario
and Quebec are poorly covered on our map due to sparse collecting in these remote areas.
 north america 57

Map NA-39. Picea engelmannii ssp. engelmannii (black)

[n=122, m=115, h=14, 1860–2008]; P. engelmannii ssp.
mexicana (red) [n=11, m=11, h=8, 1958–1994]

Picea engelmannii has an extensive distribution in the

Rocky Mountains and in the Cascade Range. Isolated
populations in the Chiricahua Mountains of SE Ari-
zona, USA and in S Chihuahua, Coahuila and Nuevo
León in Mexico are recognized as subspecies (by some
as a species) mexicana. As observed in the introduc-
tion to the genus in North America it is peculiar that
P. engelmanii does not occur in the Sierra Nevada of
California, given its far more southerly extension into Mexico. It is present in the Klamath and
Siskiyou Mountains of S Oregon and N California but absent on nearby Mt. Shasta (4322 m).
Although in Colorado and New Mexico it can reach 3600–3700 m or as very low ‘krummholz’
even higher, in N California the herbarium specimens mapped here were taken between 1050 and
1750 m while in British Columbia it can be found as low as 600 m. There is therefore no lowland
barrier preventing P. engelmannii from reaching the Sierras and it is difficult to explain, given the
fact that several other conifers with distributions in the Cascades at similar altitudes extend S into
the Sierras, what the causes are for the absence of spruces in the Sierras. It may come down to
a contingency resulting from repeated advances and retreats of tree species during the strongly
oscillating climate of the Pleistocene. Picea engelmannii forms extensive pure forests in the Rocky
Mountains, or is a constituent of mixed conifer forest especially in the Cascade Range. At the tree
line it can be on its own or is associated with Abies lasiocarpa and Pinus flexilis.

Map NA-40. Picea rubens [n=91, m=89, h=11, 1872–2010]

In apparent opposition to Picea engelmannii in the W,

P. rubens is widely distributed on the E side of North
America. In Canada, it occurs in Nova Scotia, Prince
Edward Island and New Brunswick, as well as in the
far S of Quebec and Ontario. In the USA it is distrib-
uted from Maine to North Carolina. In the maritime
provinces of Canada and in Maine P. rubens is often
found at low altitudes near the coast, but further S
it retreats into the hills and mountains of the interior until in the Southern Appalachians it is
only found on the highest ridges and summits between 1450 m and 1850 m a.s.l. It is climatically
restricted to areas with a cool, moist oceanic climate, with annual precipitation between 875 mm
and 2000 mm (increasing with elevation). In Canada and most of the New England states in the
USA this species is often associated with P. glauca; similarities between these two species can be
a source of error in the data if specimens were wrongly identified.

Map NA-41. Picea sitchensis [n=103, m=102, h=19,

1863–2005]

The well-known Sitka spruce is a coastal spe-

cies, occurring from the Semidi Islands S of the
Alaska Peninsula along the Pacific coast and
islands to Mendocino Co. in California. It is the
dominant conifer in the Alexander Archipelago
of Alaska, the Queen Charlotte Islands, Vancou-
ver Island, the mainland coast of British Colum-
bia and the coasts of Washington and Oregon,
58 north america

while in California it is replaced in this role by Sequoia sempervirens. Sitka spruce is the tallest
in its genus, with trees on Vancouver Island and in Olympic National Park, Washington reach-
ing 85–90 m tall and 4.5–5 m d.b.h. Foresters are exceedingly keen on this tree with its massive
straight bole and high quality timber; as a result big trees now mainly exist in protected areas.
In Alaska some of the large islands still have pristine coastal rain forest, but the trees while large
do not attain giant proportions this far N along the coast. Immensely important for a healthy
coastal ecosystem, it is to be hoped that Canada will follow the USA in protecting more of its
‘old growth’ forest of Sitka spruce before it is all logged. Between the mouth of Queen Charlotte
Strait at 51° N and the mouth of the Skeena River at 54° N the coast of British Columbia has no
dots on our map. Picea sitchensis is abundant there, it is just the case that botanical collecting
along this roadless coast has been piecemeal and very few conifer specimens are present in the
herbaria from this region.

Map NA-42. Picea pungens [n=94, m=93, h=6, 1862–

2006]

Picea pungens is a species of the Rocky Moun-

tains in the USA It occurs in SE Idaho, Wyo-
ming, Colorado, New Mexico and parts of Utah
and Nevada. It is most widespread in Wyoming,
Colorado and N New Mexico. It is a subalpine
species occurring in the Rocky Mountains at
elevations between 1800 m and 3300 m a.s.l.,
commonly along mountain streams or on moist,
N-facing slopes. This species grows in small, scat-
tered groves, especially near perennial streams, or scattered and mixed with Pinus con torta, P.
ponderosa, Pseudotsuga menziesii var. glauca, or Populus tremuloides. It is nearly everywhere an
uncommon constituent of the subalpine forest, but sometimes occurs outside it along streams
between more arid slopes covered in sage brush (Seriphidium tridentatum). The distribution of
Picea pungens in the ‘Four Corner States’ Utah, Colorado, New Mexico and Arizona is similar
to that of P. engelmannii in the same region (map NA-39 on p. 57) and indicates the limitations
imposed on the occurrence of spruces by climate. The Four Corners Area itself, around where
the state boundaries meet, has no spruces despite the fact that some isolated mountains e.g. the
Henry Mts. and La Sal Mts. have substantial tall conifer forests.

Map NA-43. Picea breweriana [n=41, m=37, h=14, 1885–1998]

Picea breweriana has a limited distribution in S Oregon and

N California, where it occurs in the Klamath Mountains
and in the Siskiyou Mountains. This geologically complex
region forms a transition between the volcanic Cascade
Range to the N and the uplifted granitic bloc/fault system
of the Sierra Nevada of California. It has itself high isolated
volcanoes (Mt. Shasta, Lassen Peak) of relatively young age
but also extensive areas of metamorphosed rock such as
serpentine, which is ultramafic and known for higher levels
of endemism in plants. As noted above, this region has an
exceptionally high diversity of conifers; Picea breweriana is
one of its endemics. It occurs chiefly in the Transition Life
Zone at altitudes between 1200 m and 2300 m, with e.g. Abies magnifica, A. concolor, Pseudotsuga
menziesii, Pinus monticola, P. lambertiana, P. ponderosa, Picea engelmannii, Tsuga mertensiana,
Calocedrus decurrens, Chamaecyparis lawsoniana and many shrubs. Picea breweriana is listed as
Vulnerable (VU) on the IUCN Red List.
 north america 59

Pseudotsuga

Pseudotsuga is a small genus with only two species in North America, both confined to the W of
the continent. One is widespread and extends from W Canada to Mexico; the other is a narrow
Californian endemic. Two further species occur in E Asia, one in Central and SE China includ-
ing Taiwan and N Vietnam, the other in Japan. More species have been described, on limited
evidence, and only two further taxa are here recognized as distinct at the rank of variety, one in
North America, the other in Asia.

Map NA-44. The distribution of the genus Pseudotsuga is confined to two major regions: western North
America and eastern Asia.

The past distribution of Pseudotsuga, as indicated by the fossil record, is not substantially dif-
ferent from the present. Increasing aridity has eliminated the genus from the Great Basin in the
USA, where it was present in Oligocene and Early Miocene palaeofloras. During the Pliocene it
was more widespread in Japan than it is now. This could mean that Pseudotsuga is a North Pacific
coast/Beringian genus which spread southward on either side of the ocean into its present areas,
with so far modest diversification. Phylogenetic evidence is tending towards two separate clades,
one with the Asian and one with the North American taxa (Gernandt & Liston, 1999) which
makes it difficult to determine whether a nearest common ancestor was either native to Asia or
to North America. Fossils older than the Pliocene may yet be discovered in Asia.

Map NA-45. Pseudotsuga menziesii var. menziesii

(black) [n=94, m=91, h=22, 1825–2010]; P. menziesii var.
glauca (red) [n=173, m=161, h=21, 1839–2008]

Pseudotsuga menziesii has the widest distribution

of all western North American conifers. It has
one of the greatest latitudinal ranges of any spe-
cies of conifer, extending from ca. 55° N in British
Columbia, Canada to 16° 51 N in Oaxaca, Mexico,
over a distance of nearly 5000 km. The species
divides morphologically and geographically in
two distinct varieties, a western Pacific taxon,
var. menziesii and an eastern (Rocky Mountain)
taxon, var. glauca. Our map is incomplete for
the most northern occurrences of both varieties,
in particular along the BC coast between 51° n
and 54° N, due to lack of herbarium based data.
60 north america

Whereas both varieties occur at about the same latitude at their northern limits, to the south var.
menziesii only reaches the Purisima Hills in Santa Barbara Co., California at 34° 44’ N, an isolated
population. Pseudotsuga menziesii var. glauca also becomes very scattered in Oaxaca, with only
two localities known so far. This species, known as Douglas fir, is one of the major timber trees in
North America and has been heavily exploited. There is evidence that it may have been the tallest
tree in the world in historical times (Van Pelt, 2001) but the largest/tallest trees of this species are
now ca. 100 m with 4.5–5 m d.b.h., just a few meters short of Sequoia sempervirens. These giant
trees are/were P. menziesii var. menziesii on Vancouver Island and on the Olympic Peninsula of
Washington and historically occurred all around the inland waterways opening out into the Strait
of Juan de Fuca, and on the islands. On the coast as well as in the Rocky Mountains P. menziesii
occurs mostly in mixed conifer forests; the coastal variety from near sea level to 1950 m a.s.l. and
var. glauca from 250 m in the N to 3350 m in the S of its wide range.

Pseudotsuga macrocarpa in the Sierra Madre Moun-

Map NA-46. Pseudotsuga macrocarpa [n=36, m=35, tains, Santa Barbara Co., California. Photo credit Aljos
h=11, 1867–2010] Farjon.

Pseudotsuga macrocarpa has in comparison with P. menziesii a very limited distribution; it is a

California endemic species occurring in the southern mountains from Zaca Peak in the San Rafael
Mountains in Santa Barbara Co. to Banner Grade (southernmost dot on map) and Chariot Canyon
in San Diego Co. This species is a tree of moderate size but with large seed cones. It occurs in
scattered open stands or small groves on the seaward slopes of coastal mountains at elevations
between 275 m and 2450 m, i.e. from the chaparral zone up into mixed conifer forest. Pseudotsuga
macrocarpa is not well adapted to forest fires and as fire frequency has increased due to human
causes, protection from forest fires has become a major concern. This situation is also difficult
to manage because too zealous fire fighting will build up a fuel load in these no longer exploited
stands of conifers and the inevitable fire that will occur in future may run out of control and
destroy more of this species than would have happened under natural conditions. To what extent
climate change may further complicate things is difficult to predict. Pseudotsuga macrocarpa has
been listed as Near Threatened (NT) on the IUCN Red List because management of fires has so
far been able to prevent substantial decline.

Larix

The genus Larix has 11 species, of which three are native in North America. One of these, L. laricina,
occurs across much of the northern boreal forest; the other two are more narrowly distributed in
NW USA and adjacent parts of Alberta and British Columbia in Canada. The genus is mapped and
discussed in the chapter Mainland Asia and Japan, where most species occur.
 north america 61

Map NA-47. Larix laricina [n=170, m=167, h=19, 17??–2009]

Larix laricina is one of the boreal conifers of North America with a continent-wide distribution.
Unlike Picea mariana (map NA-37 on p. 55) and P. glauca (map NA-38 on p. 56) the population in
Alaska is not connected with the main distribution of the species in Canada; there is a gap across
easternmost Alaska and most of the Yukon Territory. The Alaska population has been described
as a distinct taxon but this is not widely accepted. Like the two spruces, Larix laricina in Canada
extends to the Arctic tree line. Collecting for herbaria has been limited especially in Saskatch-
ewan, N Ontario and N Quebec, making the map incomplete; the apparent gaps there are not
real. In the E it occurs substantially further S than in the W, similar to other boreal conifers and
likewise caused by climate differences, associated with the low temperature of surface water of
the N Atlantic Ocean in winter. The northernmost locality is at Dolomite Lake in the Northwest
Territories at 68° 18’ N and the southernmost locality is in Preston Co., West Virginia at 39° 33’
N. In NE Quebec this species only reaches 58° 35’ N, almost 10° further south than in the western
part of its range. Larix laricina is a lowland species but in the SE of its range may occur to an
altitude of ca. 1000 m. It is most frequently found in swampland and muskegs where tree growth
is often sparse and stunted. Here it is most often associated with Pinus banksiana and Picea mari­
ana, on drier ground with Picea glauca, Abies balsamea and poplars and willows.

Map NA-48. Larix lyallii [n=63, m=55, h=13,

1860–2008]

Larix lyallii is a high montane to subalpine

species in the NW Rocky Mountains and
the North Cascades Mountains, occurring
on both sides of the Canada-US border. It
occurs in the Rocky Mountains of Alberta,
British Columbia, Idaho and Montana and
in a separate population 200 km to the W
in the North Cascades in British Columbia
and Washington. Its N-S extent is from 51° 37’ N near Lake Louise in Banff National Park to 45°
28’ N in the Salmon River Mountains of central Idaho. There is an eastern outlier in Montana
on Big Baldy Mountain, along Johnston Creek (alt. 2910 m) collected in 1930 (RM). Unlike the
other two American species, this larch occurs in remote, inhospitable localities near or at the
tree line, often in cirques and on scree slopes. Its northern limit is determined by the extent of
the Columbia Icefield; sub-fossil wood of probably this species growing between 1000 and 1250 CE
(the warm period of the Medieval Climate Optimum) found near the Athabasca Glacier indicates
that in historical times it grew at least 90 km further NW than now. The glaciers and ice fields
expanded again during the Little Ice Age (ca. 1550 to 1850 CE) and have been retreating since at
least ca. 1900 CE. It will take much time for this slow growing larch to follow, if indeed it will and
if, as is expected, the warming continues.
62 north america

Map NA-49. Larix occidentalis [n=70, m=68,

h=11, 1860–2005]

Larix occidentalis is widespread in the

Upper Columbia River Basin of NW Mon-
tana, N Idaho, NE Washington and SE
British Columbia. Another area of fre-
quent occurrence is formed by the eastern
slopes of the Cascade Range in southern-
most British Columbia, Washington and
N Oregon, and finally it occurs in the Blue
Mountains and Wallowa Mountains of SE
Washington and NE Oregon. This pattern
of distribution leaves a large west-central area of the extent of occurrence (EOO) without this spe-
cies. This is the Central Columbia River Basin, largely filled with extensive lava flows and lying in
the rain shadow of the Cascades. Larix occidentalis is a montane species occupying a middle zone
between low precipitation and low temperatures, from 600 m to 2100 m altitude depending on
location and exposure. In certain parts of its range it is replaced at high altitudes by L. lyallii (map
NA-48 on p. 61). Larix occidentalis can form pure stands but, depending on the successional stage,
is more often a component of mixed conifer forest. Pure stands will be invaded by more shade
tolerant evergreen conifers, which in the absence of canopy disturbance will become dominant.

Tsuga

The genus Tsuga has nine species, of which four are native to North America and five are Asian.
In North America there is a division between E and W with two species in each part of the conti-
nent. The distance between the two areas of occurrence of this genus is considerable, extending
from the Idaho-Montana border in the W to E Minnesota in the E, a distance of ca. 1700 km. Tsuga
is not a genus of boreal forest conifers although on both sides of the continent it extends into
the southern fringes of the boreal conifer forest and along the coast of the Pacific Ocean reaches
61° N near Anchorage, Alaska. In the west, the species are constituents of conifer forests, but in
the east they occur in conifer forest and in deciduous angiosperm forest.

Map NA-50. Tsuga heterophylla [n=140, m=139, h=19, 1865–2009]

 north america 63

Tsuga heterophylla has an extensive ditribution in the maritime NW of North America, where
it occurs along the coast of the Pacific Ocean from Kodiak Island in Alaska to Mendocino Co.
in California. This species is abundant along the Alaskan coast, particularly in the Alexander
Archipelago where it is the most common conifer from the high tide line into the interior of
the islands at lower altitudes. It is also abundant along the coast of British Columbia and on
the Queen Charlotte Islands and on Vancouver Island, thence S along the coast of Washington,
Oregon and N California. In N British Columbia Tsuga heterophylla starts to appear sporadically
in mountains of the interior, becoming more frequent in the Selkirk Mountains and abundant in
N Idaho and NW Montana. In Washington and Oregon it is also frequent in the Cascade Range.
Tsuga heterophylla is a dominant tree of the Pacific temperate rain forest, predominantly nearest
the coast and in river valleys. It is very shade tolerant and germinates en masse on ‘nurse logs’
i.e., large fallen trees of Picea sitchensis and Pseudotsuga menziesii. In the absence of canopy dis-
turbances (fire, storm, landslides) it would completely dominate the forest. In the interior, where
rainfall is less high, it remains only a component of mixed conifer forest, often in a second canopy
layer. At its southermost localities it is confined to streams and cool N slopes of ravines.

Map NA-51. Tsuga mertensiana ssp. mertensiana Tsuga mertensiana ssp. mertensiana in North
(black) [n=113, m=110, h=20, 1880–2006]; T. mer­ Cascades National Park, Washington, USA. Photo
tensiana ssp. grandicona (red) [n=24, m=22, h=10, credit Aljos Farjon.
1876–1992]

Tsuga mertensiana has a distribution similar to T. heterophylla (map NA-50 on p. 62) in Alaska
and Canada, but further S in the USA the two species diverge and T. mertensiana leaves the
coast and becomes a montane species, finally ending as a subalpine tree in the Sierra Nevada
of California. There its morphological distinction is recognized as a subspecies, T. mertensiana
ssp. grandicona. Also similar to T. heterophylla (and other conifer species of this region), in SE
British Columbia and adjacent parts of Idaho and Montana there is an inland Rocky Mountain
population. From a disjunct occurrence in the Chugach Mountains near Anchorage, the Alaskan
populations are in coastal mountains and islands all the way down to the Queen Charlotte Islands
and opposite parts of the Coast Mountains in British Columbia. From there, for a distance of
700 km to the southern part of Vancouver Island and the Coast Mountains opposite, there are no
herbarium based records. This is not a true gap as T. mertensiana occurs there according to USDA
Forest Survey maps and other sources. As has been observed for Picea sitchensis (map NA-41 on
p. 57) this roadless coast has been rarely visited by botanists collecting for institutional herbaria.
The same applies to the coast between Prince William Sound and Glacier Bay, but here scarcity
of this and other conifers is partly genuine as massive glaciers such as Malaspina and Bering come
near the coast and create an arctic landscape devoid of trees. Tsuga mertensiana is a subalpine
species, occurring from near sea level in Alaska to 1500 m a.s.l. along the coast; in the Cascade
Range between 1200 m and 2100 m, and in the Sierra Nevada (ssp. grandicona) between 1800 m
and 3350 m. Tsuga mertensiana is remarkable for its resilience to heavy snow packs, which in the
northerly coastal mountains can be so deep as to cover trees 15–20 m tall entirely.
64 north america

Map NA-52. Tsuga canadensis [n=109,

m=107, h=13, 17??–2009]

In the eastern half of North America,

Tsuga canadensis is the species with
a wide distribution, but unlike its
western vicariant, T. heterophylla, it is
not a constituent of extensive conifer
forests. This species occurs mainly in
a broad zone of deciduous, cool tem-
perate angiosperm (‘broad-leaved’)
forest, from Nova Scotia in the E to
Minnesota in the W. It extends southward along the entire chain of the Appalachian Mountains
to the Cumberland Plateau in N Alabama. In the maritime provinces of Canada and in Maine and
Vermont conifers may dominate; there it grows together with Pinus strobus, Picea rubens, Abies
balsamea and Betula or Populus or further S Acer and Quercus. However, T. canadensis is better
able to persist in mixed forest than most other conifers because of its relative shade tolerance.
Thin soils over bedrock, resulting from extensive glaciation only 10,000 years ago, are better habi-
tat for conifers than for most angiosperm trees and in these areas T. canadensis can form dense
stands excluding other conifer trees. Soils are acidic with pH values of 3–4 and with peat forming
in water-logged places, eventually killing even these acid-tolerant conifers.

Map NA-53. Tsuga caroliniana [n=20, m=19, h=11,

1858–1993]

Tsuga caroliniana is an endemic of the Appa-

lachian Mountains and occurs in the states of
Georgia, North Carolina, South Carolina, Ten-
nessee and Virginia. It is a relatively rare spe-
cies, restricted to cool, N- or E-facing steep rocky
slopes or along streams in ravines where it can
survive under canopy of other trees or some-
times form small pure stands. Its altitude range
is 600–1500 m, with most stands occurring between 750 m and 1200 m. This species is considered
to be a relict from the colder climate which prevailed for long periods in the Pleistocene during
glacial maxima, which were repeatedly interrupted by warmer interglacial periods, when angio-
sperms returned. However, the much more widespread and common species T. canadensis also
occurs in the Appalachian Mountains at similar altitudes but not generally in the same localities.

Abies

The genus Abies is the second most diverse genus (after Pinus) in the family Pinaceae with 47 species,
10 of which are native in North America N of Mexico. There is a division between E and W in the
distribution of the species, but in the northern boreal forest Abies balsamea meets the western spe-
cies Abies lasiocarpa in Alberta, Canada. Only one other species, A. fraseri, occurs in the E of North
America, eight species are western in their distribution, with the main diversity concentrated in the
Cascade Range and Klamath Mountains. Two species have limited ranges, one in the Santa Lucia
Mountains of California (A. bracteata) and one in the Appalachian Mountains (A. fraseri). While in
the E Abies extends no further S than the Appalachian Mountains in North Carolina, in the W the
genus continues across the US-Mexican border through Mexico to Guatemala and Honduras. Both
in terms of extent of occurrence and of taxonomic diversity the genus Abies in North America is pre-
dominantly western. Although the distribution maps of Picea (map NA-36 on p. 55) and Abies (map
NA-54 on p. 65) are similar, Abies extends much further S in California (and in Mexico). At least some
species can cope with hotter and drier summers than any in the genus Picea.
 north america 65

Map NA-54. The distribution of the genus Abies in North America (N of Mexico).

Map NA-55. Abies balsamea [n=119, m=116,

h=15, 17??–2009]

Abies balsamea is the fir of the north-

ern boreal forest, occurring from New-
foundland in the E to Alberta in the W
and extending through the New Eng-
land states as far S as West Virginia in
the Appalachian Mountains. It is par-
ticularly widespread in E Canada, New
England and the region of the Great Lakes as well as in N-central Quebec and Ontario and around
Lake Winnipeg in Manitoba. Further W in the boreal forest it becomes more scattered and its most
western localities are E of the Rocky Mountains in W-central Alberta. As with other boreal conifers,
a lack of herbarium collections from these vast territories (we found none from Saskatchewan)
makes the map incomplete. In the W this species meets A. lasiocarpa (map NA-56) but that species
is predominantly subalpine while A. balsamea is a lowland species which only occurs at higher alti-
tudes (1200–1600 m) at its southernmost stations in the Appalachian Mountains (up to 1900 m on
Mt. Washington in New Hampshire, where it becomes prostrate due to severe winter storms). Abies
balsamea is a constituent of coniferous forests with Picea spp., Pinus strobus, Tsuga canadensis and
sometimes Pinus banksiana, or it grows mixed with angiosperm trees such as Populus tremuloides,
Betula spp. and, further south, Acer spp., Fagus grandifolia and Betula alleghaniensis.

Map NA-56. Abies lasiocarpa var. Abies lasiocarpa var. lasiocarpa in Mount Rainier National Park,
lasiocarpa (black) [n=143, m=141, h=17, Washington, USA. Photo credit Aljos Farjon.
1831–2010]; A. lasiocarpa var. arizonica
(red) [n=16, m=16, h=7, 1908–2004].
66 north america

The distribution of Abies lasiocarpa is for the most part in the Rocky Mountains, extending from
the Northwest Territories and Yukon Territory in the N to Arizona and New Mexico in the S.
It is also present in the Coast Mountains and islands of S Alaska and British Columbia, on the
Olympic Peninsula in NW Washington, in the Cascade Range, including its high volcanoes, and
in the Blue Mountains of NE Oregon, terminating in the Klamath Mountains in N California. In
the southernmost section of the Rocky Mountains A. lasiocarpa var. arizonica occurs in the ‘Four
Corner States’, but of these mostly in Arizona and New Mexico. The typical habitat of this spe-
cies is the subalpine zone between closed conifer forest and treeless alpine meadows. This zone
occurs on average only a few hundred meters a.s.l. in the northernmost part of its range and rises
to around 3000 m in the farthest south. It is generally lower in the Coast Mountains and the Cas-
cade Range, which receive higher levels of precipitation (especially snow) than most of the Rocky
Mountains. The two conifers most often associated with A. lasiocarpa are Tsuga mertensiana in
the coastal mountains (map NA-51 on p. 63) and Picea engelmannii in the Rocky Mountains (map
NA-39 on p. 57).

Map NA-57. Abies grandis [n=94, m=93, h=19, 1857–2006]

Abies grandis is distributed in the temperate coni-

fer rain forest of the Pacific Northwest and occurs
abundantly from Vancouver Island S to Sonoma
Co. in California. Its northernmost locality is near
Prince Rupert in British Columbia, but this far N
it is uncommon even though the collection bias
mentioned earlier (e.g. Picea sitchensis map NA-41
on p. 57) gives too few dots on our map. In the lati-
tudinal middle part of its range between ca. 44° N
and 49° N A. grandis extends eastwards from the
Cascade Range to the Blue Mountains in Oregon
and to the Rocky Mountains around Kootenay
Lake in BC and in Idaho and, more sporadically, in
W Montana. These eastern extensions are separated from the main western range by the Columbia
Plateau, an extensive area of ancient lava flows cut through by the Columbia River and its tribu-
taries. Sufficient precipitation at this latitude picks up again on the mountains immediately E of
this plateau and several rain forest conifers ‘reappear’ there (compare the maps of Taxus brevifolia
NA-91 on p. 87, Tsuga heterophylla NA-50 on p. 62, T. mertensiana NA-51 on p. 63 and Thuja plicata
NA-69 on p. 74). Or, perhaps, they disappeared from the intervening region when the Cascades
were elevated to sufficient height to block much of the rain coming from the Pacific Ocean.

Map NA-58. Abies procera [n=31, m=30, h=8, 1825–2005]

Abies procera occurs in the states of Washington and

Oregon in the W USA, where it is common in the Cas-
cade Range from Stevens Pass in King Co., Washing-
ton to Grasshopper Mountain and vicinity in Lane Co,
Oregon. There are also a few localities with this species
nearer the Oregon coast. This species is absent in SE
Oregon where it is replaced by A. magnifica var. shas­
tensis (map NA-59 on p. 67) and according to some,
intermediate hybrid forms between these taxa. These
seem to occur within a broad zone with its southern
limit around Lassen Peak in N California (R. J. Laacke
in Burns & Honkala, 1990). We have mapped only
unambiguously identified herbarium specimens of
 north america 67

A. procera here; if introgression via pollen from A. procera into A. magnifica does indeed occur,
as experiments indicate, the S limit of A. procera may not be as abrupt as our map suggests.
Specimens in herbaria identified as A. procera and collected in California belong most likely to
A. magnifica var. shastensis; A. procera has not been found in that state (Debreczy & Rácz, 2011,
vol. 2: 583).

Map NA-59. Abies magnifica var. magnifica (black) [n=59,

m=56, h=17, 1853–2003]; A. magnifica var. shastensis (red)
[n=38, m=38, h=13, 1852–2006]

Abies magnifica is mainly a Californian species which

extends N into Oregon along the Cascade Range. It
occurs along the length of the Sierra Nevada and there
is an isolated population in the San Bernardino Moun-
tains at 34° 20’ S. In N California it is common in the
Klamath, Siskiyou and Coast Mountains, and here the
variety shastensis (named after Mt. Shasta) is also pres-
ent, occurring to the N as far as Crater Lake in Oregon.
Another outlying population of A. magnifica is found
in the Warner Mountains of extreme NE California and
near Lake Tahoe this species crosses the border into
Nevada. The two varieties are largely sympatric in NW
California but in some areas var. shastensis dominates.
Seed cones with clearly exserted bracts (a character
state of var. shastensis) also appear on trees of A. magnifica in the S part of the Sierra Nevada in
Madera, Fresno and Tulare Counties. If these are also A. magnifica var. shastensis there would be a
ca. 400 km gap between the two populations of this variety. The identity of these southern trees is
not certain and needs verification. Abies magnifica is a tall growing fir growing with other conifers
in mixed conifer forest, sometimes forming pure stands and in other places scattered among dif-
ferent species. Its altitude range is generally between 1400 m and 2700 m corresponding with the
Canadian Life Zone of the W slopes of the Sierras, one of the richest and most magnificent conifer
forests in the world.

Abies magnifica in the Sierra Nevada above Tamarack

Creek, Fresno Co., California. This stand of mature trees
is in the Sierra National Forest and has been selectively
logged; abundant regeneration can be seen in the area
behind the tall trees. Similar open spaces, colonized by
conifers from seed dispersed from trees left standing,
would be created by storms or fires in totally unmanaged
forests. In national parks, such as Kings Canyon, Sequoia
and Yosemite N.P. in these mountains, no trees are
felled, but fires were traditionally prevented or put out
if possible. Slowly, the more shade tolerant conifers are
taking over many areas and regeneration of this kind
is becoming uncommon, or it might in future occur on
a large scale when fires have become unmanageable
due to build up of fuel load. Perhaps selective logging
is not so bad for forest diversity as the bad press it has
received from over-zealous conservationists suggests.
Photo credit Aljos Farjon.
68 north america

Map NA-60. Abies amabilis [n=67, m=63, h=12, 1825–

2002]

Abies amabilis is another conifer species native to

the Pacific Northwest, occurring from the Coast
Mountains in British Columbia at ca. 56° N and
extreme SE Alaska to Vancouver Island, the Olym-
pic Peninsula of Washington and the Coast and
Cascade Ranges S as far as the Klamath Moun-
tains in N California. It does not occur further
than 200–250 km from the Pacific coast but is also
absent from more oceanic islands such as in the
Queen Charlotte Group and the outer islands of S
Alaska, which are well forested. Herbarium based
data along the mainland coast of BC are largely
missing, as for other species, and the apparent gaps
on the map are artificial. Abies amabilis occurs at
low altitudes in the N of its range and ascends to
ca. 1800 m in the S. It is a constituent of mixed
conifer temperate rain forest, with annual precipitation ranging from 1500 mm to 4000 mm, near
the coast mostly as rain but at higher elevations in the Coast and Cascade Ranges also in the form
of heavy winter snow packs.

Map NA-61. Abies concolor [n=123, m=122,

h=17, 1847–2005]

Abies concolor is widely distributed in the

western USA and extends a short distance
into Mexico. The species here includes
populations in S Oregon, California and
N Baja California which are sometimes
treated as a separate taxon (A. lowiana or
A. concolor var. lowiana); in the N of this
range it may hybridize with A. grandis
(map NA-57 on p. 66) so A. lowiana has
also been described as A. grandis var. lowi­
ana. In California the main distribution of
A. concolor is in the Sierra Nevada, where it
occurs on both sides of the divide, includ-
ing a small corner of Nevada. It is also
common in N California and a short distance into Oregon. In S California it is present in most of
the highest mountains and a distant southernmost location is in the Sierra San Pedro Martír, Baja
California Norte, Mexico. Eastward, it is present in mountains of the ‘Basin and Range Province’
or Great Basin, sporadically in E Nevada and more commonly in the Wasatch Range of Utah. In
Arizona it occurs mostly S of the Colorado Plateau in the volcanic San Francisco Mountains, on
the Mogollon Rim and mountains in the SE, from where it extends S to a few localities in E Sonora
and W Chihuahua, Mexico. Greater abundance occurs in the Rocky Mountains of Colorado, but
only in the southern half of the state, and in New Mexico. Abies concolor occurs from 650 m
to 3350 m altitude and is more tolerant of dry spells and high light intensity than most other
species in the genus. It often occurs in more or less open conifer woodland in association with
other species.
 north america 69

Map NA-62. Abies fraseri [n=30, m=28, h=8,

1876–2006]

Abies fraseri has a limited distribution in

the southern Appalachian Mountains,
mainly in North Carolina but crossing
into Tennessee in Great Smoky Mountains
National Park, and with a few northern-
most localities in Virginia. Major stands
occur on Mt. Mitchell and surrounding
mountains and on Grandfather Mountain
in North Carolina. Abies fraseri is limited
to the highest mountains at altitudes
between 1200 m and 2150 m, often best developed on N-facing slopes. It is closely related to A. bal­
samea (map NA-55 on p. 65) but geographically separated from it by a distance of ca. 180 km. Acid
rain from industrial air pollution has devastated many stands, leaving thousands of dead standing
trunks, but after the clean-up of sulphur dioxide emissions a few decades ago re-establishment
from seedlings is taking place and in due course these populations may recover. Perhaps a more
threatening agent is the introduction, first discovered on Mt. Mitchell in 1957, of an alien scale
insect that has killed very large numbers of trees. At present A. fraseri is listed as Endangered
(EN) on the IUCN Red List due to this threat for which no remedy, other than using insecticides,
has yet been found effective.

Map NA-63. Abies bracteata [n=39, m=37, h=16, 1831–1997]

Abies bracteata is endemic to the Santa Lucia Moun-

tains on the coast of California S of Monterey and
occurs from the headwaters of the Carmel River and
Little Sur River along the high parts of the mountain
range to the headwaters of the Arroyo de la Cruz in
San Luis Obispo County. The southernmost locality
from where a herbarium specimen exists (Chester
Dudley s.n., 23 Jul 1927, CAS 147657) is (upper) Las
Tablas Creek, San Luis Obispo. The great majority of
stands are in Monterey County. Abies bracteata occurs
in canyons and on steep slopes and ridges at the head-
waters of numerous creeks and small rivers above a
zone with Sequoia sempervirens in mixed evergreen forest. It is an emergent above the canopy
of this forest, or it occurs on rocky ridges and summits where other trees cannot grow well. It is
never further than ca. 20 km from the coast; the two most inland localities are Anastasia Canyon
and Bear Canyon. Its altitude range is 600–1570 m but in deep canyons a few stands are known
at lower elevation. Abies bracteata is an unusual fir that is not closely related to any other species
in the genus and has been found to be ‘basal’ to all species in some cladistic analyses based on
molecular data. It may be a representative of an early-diverging lineage of the genus now con-
fined to this small mountain range on the Pacific coast.
70 north america

Abies bracteata in a ravine on the upper slopes

of the Santa Lucia Mountains, Monterey Co.,
California. This species has long been classified
in its own section or subgenus within the genus
Abies on account of several unique morphological
characters. This distinct position has been confirmed
by more recent molecular (DNA) evidence and in
phylogenetic analyses using DNA sequence data this
species usually appears as a basal clade in relation
to the other species in the genus. Is it perhaps a
primitive relict species? The fossil record does not
help much here, as fossilized remains of the genus
Abies are scarce and nothing has been found that
resembles this species. The timing of the split of that
basal clade from the rest of the genus by ‘relaxed
molecular clock’ calculations is hampered by the
same lack of well dated fossils, used to calibrate the
clock. The genus is surely much older than the oldest
fossil found and most probably the closest relatives
of this peculiar fir became extinct a long time ago.
They are the last survivors of an ancient lineage.
Photo credit Aljos Farjon.

Cupressaceae

The Cupressaceae are the second most prominent family of conifers in North America, in some
cases forest-forming but more commonly occurring mixed with conifers in the Pinaceae or less
often mixed with angiosperm trees and shrubs. These forests range from the extremely wet Pacific
Northwest temperate evergreen rain forest to juniper woodland in the dry, semi-desert interior
of the continent, and from the subarctic tree line in the northern boreal forest to the cypress
swamps of Florida and the Gulf of Mexico. This range of habitats across the continent is similar
to that occupied by Pinaceae, but extends to greater extremes both on the wet and on the dry
ends of the climatic and edaphic amplitudes. This is to some extent reflected in the distribution
of the family within the contiguous USA (map NA-2 on p. 36) where species in the Cupressaceae
occur in parts of the continent not occupied by members of the Pinaceae.

Sequoioideae

This subfamily comprises three genera (Farjon, 2005a), two of which occur in North America:
Sequoia and Sequoiadendron (the third, Metasequoia, is confined to a small area in China). This
small group (clade) of three monospecific genera is remarkable because the current distribu-
tion of its species is but a fraction of the extent in the geological past. All three species were
widespread across the Northern Hemisphere in the Tertiary up to the Miocene or Pliocene, after
which time they retreated further and further until they ended up in their present refuges. The
cold climate of the Pleistocene greatly hastened their demise. The next ‘ice age’ could be the
cause of their extinction if they had remained in these limited locations, but they have now been
planted widely in nearly all continents. The two American species are also remarkable both for
their size and longevity; they are the tallest and largest trees in the world and can live beyond
2000–3000 years.
 north america 71

Sequoia sempervirens at Bull Creek, Map NA-64. Sequoia sempervirens [n=83,

Humboldt Co., California. Photo credit m=76, h=14, 1794–1997]
Robert Van Pelt.

Sequoia sempervirens is a coastal species endemic to the California Floristic Province; it occurs
along the Pacific coast from the Chetco River in Curry Co., Oregon at 42° 11’ N to the border
between Monterey and San Luis Obispo Counties in California at ca. 35° 45’ N. Its furthest local-
ity inland is probably Wooden Valley in Napa Co., NE of the San Francisco Bay Area. It can
occur very close to the ocean, immediately behind a narrow strip of dunes or even right up to
the beach in some northern localities. Here it dominates the forest and trees over 100 m tall are
not rare, with the tallest tree found measuring 115 m. Further inland Sequoia sempervirens can
occur to 920 m a.s.l. and here it is more commonly mixed with other conifers, e.g. Pseudotsuga
menziesii, Tsuga heterophylla and Abies grandis. On the coast it thrives within the fog belt, a zone
only a few km wide in many places where daily fog brings moisture and reduces heat from the
sun; inland localities are mostly following streams. Sequoia sempervirens regenerates (apart from
seed) by sending out suckers from a massive lignotuber; after a destructive fire one or more of
these may develop into new trees. In the fossil record Sequoia is abundantly present from the Late
Cretaceous to the Pliocene in W North America and Europe (Florin, 1963). In the Late Paleocene
and Eocene it may have occurred in the Arctic, with fossil records in Greenland and Spitsbergen,
and in the subarctic Alaska Peninsula. Not all these fossils can unambiguously be assigned to
Sequoia. Both the Beringian migration route and one via Greenland were available at this time
for a genus of which the earliest fossils known are North American. More scattered fossil finds
dated Oligocene to Miocene are from Central Asia and the Altai Mountains. In the Pliocene it still
existed in W USA, Europe and Japan; towards the end of that period 2.5 Ma Sequoia disappeared
from Eurasia and contracted in North America, retreating further during the Pleistocene to its
present narrow coastal range almost entirely in the northern half of California. Massive logging,
carried on until recently, has caused substantial reductions in the total area of occupancy (AOO),
but the extent of occurrence (EOO) as shown on the map has remained largely the same as before
European settlement. This species is listed as Endangered (EN) on the IUCN Red List.
72 north america

Map NA-65. Sequoiadendron giganteum [n=43, m=42, h=14,

1853–2004]

Sequoiadendron giganteum is confined to the Sierra

Nevada of California, occurring in Calaveras, Fresno,
Madera, Mariposa, Placer, Tulare and Tuolumne
Counties. Its past distribution is less clear than that
of Sequoia due to difficulties with identifying its fos-
sils unambiguously, but the genus (or close relatives)
seems to have been present in North America and
in Europe since the Upper Cretaceous (Stockey et al.
in Farjon, 2005a). It is known to occur now in ca. 67
‘groves’ which stretch over a length of 400 km along the
W slopes of the Sierras in a belt at most 25 km wide,
covering a total area of occupancy of ca. 14,200 ha.
Most of the groves (59) are in the two southernmost
counties: Fresno and Tulare. There may be as few as 10 or as many as 20,000 individuals in a
grove, counting all age classes. Although there may be small stands excluding other trees, in most
localities S. giganteum occurs in association with other conifers, among which Abies concolor and
A. magnifica as well as Pinus lambertiana are the most prominent. These groves have their origin
in massive regeneration after a major forest disturbance, a phenomenon which can be observed
in progress in the Converse Basin Grove, Fresno Co., one of the largest stands, which was com-
pletely logged out save for one giant tree around the turn of the 19th/20th century. In the absence
of renewed disturbance the more shade tolerant firs (Abies) become abundant and will prevent
regeneration of S. giganteum, so it may disappear despite the potential age of 3000+ years of an
individual giant tree. This process has now started because protection in the national forests and
parks where nearly all groves are situated has meant prevention of fires for more than a century.
Despite its ‘protection’ in national parks and other protected areas, S. giganteum met the criteria
and is now listed as Endangered (EN) on the IUCN Red List.

Taxodioideae

This subfamily has three genera, only one of which, Taxodium, is native to North America. Two spe-
cies are recognized, T. distichum and T. mucronatum; the former occurs in the USA and the latter
mostly in Mexico and is treated in the chapter Mexico, Central America and the West Indies (map
MCW-56 on p. 122). Taxodium was widely distributed in W North America and in Europe from the
Paleocene to the Pliocene (Florin, 1963) but became extinct in both regions at the end of the Plio-
cene when the climate became much colder, leading to the massive glaciations in the Pleistocene.
Because no fossils older than Pliocene are known from E North America and Mexico, it is possible
that Taxodium migrated from W to E in North America sometime in the later Neogene.

Map NA-66. Taxodium distichum var.

distichum (black) [n=98, m=85, h=21,
1640–2005]; T. distichum var. imbricarium
(red) [n=41, m=40, h=11, 18??–2001]

The distribution of Taxodium disti­

chum follows the Atlantic Coastal
Plain from S New York to Texas and
follows the Mississippi River and
lower reaches of its tributaries from
Indiana to New Orleans. Its habitat,
slow running lowland rivers and
adjacent bayous and swamps above
 north america 73

the reach of salt water, is extensive in these regions. This species is distributed mostly in the far
SE of the USA and along the Atlantic coast, where the climate is warm temperate, but localities in
the interior and N of the range experience frost in winter, at times severe. Taxodium distichum is
deciduous, dropping foliage branchlets before winter frost occurs, and experiences a distinct win-
ter rest period similar to deciduous larches (Larix) which occur much further N (e.g. L. laricina
map NA-47 on p. 61). Frosts are rare or non-occurring in S Florida, but the trees are deciduous
there, too. If there is an analogy with angiosperms in this deciduous habit, it may have evolved
during a time when Taxodium occurred further N and W prior the the Pliocene migration as
surmised above. Two varieties are recognized, T. distichum var. distichum and T. distichum var.
imbricarium. They are sympatric in many areas, but var. imbricarium has a more southerly distri-
bution (as far as the Florida Keys) and is absent in the northernmost states in which the species
occurs. This variety, known as Pond Cypress, tends to occur in swamps and around lakes away
from rivers, where the water is generally less eutrophic.

Cupressoideae

The remainder of genera and species in the family Cupressaceae occurring in North America belong
to this subfamily. The genera are Chamaecyparis, Cupressus, Juniperus, Thuja and Xanthocyparis,
of which Juniperus is by far the most diverse in species and the most widespread. Chamaecyparis
and Thuja, each with two species in the continent, display the same E-W disjunction we see in
Tsuga (Pinaceae) and Taxus (Taxaceae). Cupressus and the closely related genus Xanthocyparis
are exclusively western, while Juniperus is spread across the continent from E to W and N to S.
Evidently, of these genera Juniperus has been most successful in adapting to a great variety of
habitats and consequently spread, helped by bird dispersal of the seeds, and speciated. All these
genera have congeners in Eurasia, but only in the case of two, Cupressus and Juniperus, does this
include Europe.

Chamaecyparis

Map NA-67. Chamaecyparis thyoides var. thyoides

(black) [n=56, m=51, h=12, 17??-2008]; C. thyoides var.
henryae (red) [n=21, m=20, h=9, 1896–1998]

Chamaecyparis thyoides occurs on the Atlantic

Coastal Plain from New Hampshire in the N to
Louisiana in the S, with var. thyoides compris-
ing all populations N of Florida and the coast
of Alabama and Mississippi, where it is almost
entirely replaced by var. henryae. Chamaecy­
paris thyoides is particularly common in Mas-
sachusetts and New Jersey (Pine Barrens) and
in NE Florida. Both in the N and S of the species
range it occurs near the coast as well as further inland, but in North Carolina, South Carolina and
Georgia it is not present near the Atlantic Ocean but occurs about halfway between the coast
and the Piedmont. It is not apparent why this should be so, but it is notable that this species and
Taxodium distichum (map NA-66 on p. 72) are mostly allopatric outside N Florida. Chamaecyparis
thyoides replaces Taxodium distichum from the New Jersey Pine Barrens to New Hampshire while
the reverse is the case on the near-coastal plain from North Carolina to S Florida. Chamaecyparis
thyoides usually grows in more or less pure stands in bogs and swamps and along streamside cor-
ridors of lowland rivers surrounded by other tree species. It avoids salinity although it is known
to border tidal marshes in New Jersey. The water table is always high and often there is prolonged
inundation in winter. It is likely that recurring fire would historically have been the disturbance
74 north america

agent preventing Acer rubrum from eventually replacing C. thyoides in the succession. Outside the
wettest areas in swamps and woods in the NE of its range fire prevention may be a factor adding
to the effects of past timber exploitation causing decline of this species.

Map NA-68. Chamaecyparis lawsoniana [n=49, m=47,

h=16, 18??–2008]

Chamaecyparis lawsoniana occurs in a limited area

on either side of the California/Oregon border (four
counties in California and five counties in Oregon). Its
main distribution areas are the Klamath Mountains
in Oregon and the Siskiyou Mountains in California.
To the E in California there is another concentra-
tion on the W side of Mt. Shasta and vicinity and
in Oregon there are more scattered occurrences on
the western slopes of the Cascade Range and along
the coast. It occurs in mixed conifer forest or mixed
angiosperm-conifer forest on a variety of rock and
soils, including ultramafic substrates derived from
serpentine. The altitude ranges from near sea level
to 1950 m. The limited natural range of this species
is in stark contrast to its spread as an ornamental
tree, of which more than 450 registered cultivars are
known (Auders & Spicer, 2012). In contrast, the spe-
cies in its natural habitat is now being depleted by
the introduced (1952) pathogenic oomycete Phytophthora lateralis causing root rot. Logging has
not only reduced subpopulations in terms of mature trees, but opened up areas with logging
roads along which the pathogen has been transported into previously non-infected areas. Drastic
measures such as a ban on further logging, restrictions of access and research into resistant geno-
types to be used for re-introduction in devastated areas are taken to halt and perhaps reverse the
decline. This species is currently listed as Near Threatened (NT) in the IUCN Red List.

Thuja

Map NA-69. Thuja occidentalis (black) [n=120, m=111, h=11, 17??–2010]; Thuja plicata (red) [n=119, m=106,
h=18, 1794–2004]

These two species occupy different parts of the North American continent with a clear division
between E and W, similar to other genera, e.g. Tsuga and Taxus. Thuja occidentalis is widespread
in the eastern half of the continent. [The epithet occidentalis means ‘from the west’ but this is
 north america 75

because Linnaeus when he named the species knew only one other species, T. orientalis (now
Platycladus orientalis) from China; as seen from Europe T. occidentalis was in the W, and T. ori­
entalis in the E.] It occurs from New Brunswick and Nova Scotia to Manitoba in the W, and S as
far as North Carolina and Tennessee. In the boreal conifer forest it is uncommon except near the
coast, its main distribution lies in the mixed conifer-angiosperm forest zone, and to the S it is
present but not abundant in the deciduous angiosperm forest zone. This species is often found
along the margins of calcareous fens or in open woodland in rocky terrain. Thuja plicata is dis-
tributed in W North America and, like some other conifer species, occurs there in two distinct
regions. The most extensive one is in the coastal evergreen temperate rain forest from far S Alaska
and the Queen Charlotte Islands along the coast of British Columbia and Vancouver Island to
W Washington and W Oregon, ending in NW California. The smaller area of occurrence is in
the Rocky Mountains of SE British Columbia, W Montana, N Idaho and NW Washington. In the
coastal rain forest T. plicata grows to a very large tree and is associated with Tsuga heterophylla,
Picea sitchensis, Pseudotsuga menziesii and Abies grandis; in the interior it is also a constituent
of mixed conifer forest. Limits of low temperature restrict its occurrence further N in the Rocky
Mountains and limits of lower rainfall further S; in N-central Idaho most specimens were col-
lected from streamsides or lakesides.

Thuja occidentalis (middle foreground) with junipers (low shrub) and pines on Flying Mountain, Acadia
National Park, Maine, USA. Photo credit Aljos Farjon.

Cupressus

The genus Cupressus has a very scattered distribution in the Northern Hemisphere. [In this Atlas,
as in A Handbook of the World’s Conifers (Farjon, 2010a) and in Conifers around the World
(Debreczy & Rácz, 2011) we maintain Cupressus as ‘traditionally’ circumscribed. Outside the cla-
distic paradigm there are no good evolutionary arguments for a split between ‘New World’ and
‘Old World’ cypresses at genus rank and even within this concept not all published analyses agree,
depending on what molecular data and taxa are included or excluded.] In North America Cupres­
sus is exclusively SW in distribution, extending into Mexico and Central America; in Eurasia and
Africa it has two regions of occurrence, the Mediterranean and adjacent deserts and mountains
to the S and E, and the Sino-Himalayan mountain system with extension further E in China. Of a
total of 15 species seven are native in North America and one in Mexico to Honduras.
76 north america

Map NA-70. Global distribution of the genus Cupressus.

Species in this genus vary in their level of adaptation to limits of precipitation from extremely
drought tolerant to dependency on high levels of monsoonal rainfall. In the Americas, only one
species, C. lusitanica (map MCW-42 on p. 115) occurs in a mesic environment, the mountains
of Mexico and Central America. The species of North America are all adapted to xeric environ-
mental conditions. Even greater extremes occur within this genus in Africa and Eurasia, where
species vary between C. dupreziana in the Sahara (map EM-30 on p. 159) and C. cashmeriana in
the Himalaya of Bhutan (map MAJ-26 on p. 191). Although morphological adaptations to aridity
are evident through comparative analysis and physiology, phylogenetic analyses appear to indi-
cate that these traits developed independently of relationships (Little, 2006) in both the western
and eastern hemispheres. The fossil record is poor on Cupressus or at least on taxa that may be
assigned to it with confidence (Stockey et al. in Farjon, 2005a).

Map NA-71. Cupressus sargentii [n=66, m=65,

h=12, 1880–2009]

Cupressus sargentii is endemic to Cali-

fornia but is widely distributed in that
state. There are two main areas, one N
of San Francisco Bay and another in the
Santa Lucia Mountains of Monterey Co.
The northern (sub)populations are in the
southern part of the North Coast Range
and here single ‘stands’ can be exten-
sive especially in valleys of small rivers
or creeks. It occurs mostly inland in this
northern area, where C. goveniana (map
NA-74 on p. 77) replaces it along the
coast. In the S of its range it comes nearer
the coast as it is mostly restricted to the
W slopes of the Santa Lucia Mountains.
Two isolated localities are known in the
San Rafael Mountains of Santa Barbara
Co., these are the southernmost occur-
rences of this species. Cupressus sargentii is strongly associated with ultramafic soils and rocks
derived from serpentine, to the extent that the geology has a crucial influence on its distribution
(Griffin & Critchfield, 1972). This species is listed as Vulnerable (VU) on the IUCN Red List.
 north america 77

Map NA-72. Cupressus macnabiana [n=43, m=40, h=8, 1853–1972]

Cupressus macnabiana is limited to N California, where it is

partly sympatric with C. sargentii but extends further inland
and northward. The most extensive (sub)populations are in
the North Coast Range, close to some of the of C. sargen­
tii populations or they occur indeed mixed with this spe-
cies. It may react better to fires as it occurs often on higher
slopes away from streams and is usually shrubby although
it can attain large tree size. Although often on serpentine,
C. macnabiana is found on a variety of rock types in the
foothills of the Sierra Nevada and Cascade Range, including
greenstone, conglomerate and basalt.

Map NA-73. Cupressus bakeri [n=31, m=30, h=10, 1898–1998]

Cupressus bakeri occurs in N California and across the bor-

der with Oregon in Jackson Co. and Josephine Co. It is much
less common than the previous two species both in numbers
of localities, (sub)populations and individual trees. Only two
large populations are known, near Timbered Crater and on
Seiad Creek; all others are much smaller and often many km
apart. Sometimes a historically known population has not
been found again and is probably extinct, possibly from a fire
that burnt too long and too hot for any trees or their seeds
to survive. As far as known the dots on our map all represent
existing populations and include those recorded in Griffin &
Critchfield (1972) plus a new one in Oregon. Cupressus bakeri
occurs at higher altitudes than the other Californian species
at 800–2100 m and consequently more often in mixed coni-
fer forest than in oak scrubland. A seemingly contradictory
combination of too frequent fires and fire suppression, both
man-made, is preventing regeneration and has caused the
loss of subpopulations and a continuing decline. This species
is listed as Endangered (EN) on the IUCN Red List.

Map NA-74. Cupressus goveniana var. goveniana (black) [n=39,

m=39, h=10, 1848–1997]; C. goveniana var. abramsiana (red) [n=14,
m=14, h=9, 1882–1992]

Cupressus goveniana is a coastal species endemic to Cali-

fornia. It occurs from around Fort Bragg in Mendocino Co.
S to Gibson Creek in Monterey Co. in scattered localities in
three main areas. The Mendocino Co. populations (‘Men-
docino Cypress’) occur partly on raised old coastal terraces
of podzolic sands where the trees are dwarfed; these have
been named Cupressus pygmaea. These flats also support a
dwarfed form of Pinus con torta, but both trees grow much
taller where they occur in the margins of surrounding Red-
wood forest or in gullies where organic matter has enriched the soil. One location with ‘pygmy’
cypresses is 27 km inland at Old Ornbaun Hotsprings. S of San Francisco Bay C. goveniana appears
in scattered populations inland, most of these are identified as var. abramsiana (formerly at spe-
cies rank). Furthest S C. goveniana occurs on the Monterey Peninsula mainly on Huckleberry Hill
and S from there along Gibson Creek. Cupressus goveniana var. abramsiana is severely threatened
78 north america

in an urbanizing area with high risk of widldfires that are rigorously put out if possible; it is listed
as Critically Endangered (CR) on the IUCN Red List. The status of var. goveniana is somewhat less
perilous due to its wider occurrence: it is listed as Endangered (EN).

Cupressus macrocarpa during fog conditions at Point Lobos,

Monterey Co., California. Photo credit Aljos Farjon.

Map NA-75. Cupressus macrocarpa [n=18,

m=15, h=9, 1847–2002].

Cupressus macrocarpa has a very limited natural distribution in Monterey Co., California. It is most
abundant along the rocky coast of the Monterey Peninsula from Cypress Point (the most W cape) to
Pescadero Point at the N end of Carmel Bay. Scattered occurrences have been found in Del Monte
Forest and at Asilomar State Beach. At the S end of Carmel Bay a population of C. macrocarpa
occurs in Point Lobos State Park. This species is in its natural habitat confined to the narrow fog
belt along the coast and appears to occur from the high tide line to less than 1 km inland, either in
pure stands or mixed with Pinus radiata var. radiata (map NA-19 on p. 46). Yet it has been planted
in many parts of California, where it often naturalizes, and throughout the world as an ornamental
tree or, more abundantly, as plantations and wind breaks. The natural populations are remnants of
a once wider distribution, pressed to the coast where they persist more or less out of reach of fires
due to the daily fog banks coming in from the ocean. The fog is caused by the cold California Cur-
rent and the movement of warm air over land on to the cold waters of the ocean (advection fog);
it evaporates during the day from inland towards the coast. The width of the fog belt is dependent
on its depth above the ocean, with shallow depth causing a narrow fog belt. In Monterey it is often
very narrow, restricting this fire sensitive species to the vicinity of the sea. Although the species
is legally protected, encroaching urbanisation and tourism increase the risk of destructive fires;
C. macrocarpa is listed as Vulnerable (VU) on the IUCN Red List.

Map NA-76. Cupressus guadalupensis var. forbesii (black) [n=31,

m=24, h=12, 1875–2004]; C. guadalupensis var. guadalupensis
(red) [n=8, m=8, h=8, 1875–1991]

Cupressus guadalupensis and C. forbesii were described as

separate species and their distribution is certainly disjunct,
with C. guadalupensis var. guadalupensis on Mexican Gua-
dalupe Island 240 km from the coast of Baja California and
var. forbesii on the mainland N and S of the Mexico-USA
border. The (sub)populations or ‘stands’ on the mainland
are scattered, with one locality in Orange Co. (Sierra Peak
at 33° 53’ N) and a few in San Diego Co., continuing S to
 north america 79

and across the Mexican border as far as Arroyo Hediondo at 31° 14’ N. The Mexican populations
are mostly ‘hidden away’ in isolated canyons. The threat from fires is probably less great there
than in the N (California) due to remoteness. On Guadalupe Island the main threat was caused
by introduced goats but these animals have now finally been eradicated. Both varieties are listed
as Endangered (EN) on the IUCN Red List.

Map NA-77. Cupressus arizonica var.

arizonica (black) [n=57, m=53, h=14,
1848–2000]; C. arizonica var. glabra (red)
[n=20, m=20, h=8, 1903–1995]; C. arizonica
var. nevadensis (orange) [n=19, m=18, h=10,
1918–1998]; C. arizonica var. stephensonii
(purple) [n=15, m=15, h=11, 1934–1991];
C. arizonica var. montana (blue) [n=7,
m=5, h=9, 1930–1996]

Cupressus arizonica is a widespread

and variable species, occurring in the
SW USA and in Mexico. The nomi-
nate variety arizonica has the widest
distribution and occurs in Arizona, W New Mexico, SW Texas and N and central Mexico. It could
be considered a Mexican taxon with extension into the USA, but it was first described as a spe-
cies from Arizona and several of the recognized varieties occur only N of the Mexican border.
Var. glabra is restricted to an area in central Arizona mostly N of the Mogollon Rim. Younger
trees with smooth bark cannot be certainly assigned to this variety and may belong to var. ari­
zonica, as is likely the case with some records from Mexico. The three Californian varieties have
very limited distributions well separated from each other and perhaps because of this have been
described and named as separate species. Var. nevadensis occurs at the southern end of the Sierra
Nevada, mainly in Kern Co. around the town of Bodfish, but with one locality (Greenhorn Mts.)
in Tulare Co. and to the S an outlier in Los Angeles Co. on Warm Springs Mountain. Var. ste­
phensonii occurs on both sides of the USA/Mexico border; the N population is at the headwaters
of King Creek in San Diego Co. and the S population in the Sierra de Juarez in Arroyo Seco and
(La) Rincón. Finally, var. montana occurs in several localities in the Sierra San Pedro Martír, the
highest mountain complex in Baja California. Most of these Californian varieties form ‘groves’
or (sub)populations of trees of tens to thousands of individuals associated with few other trees,
but var. montana occurs at higher altitudes in the zone of mixed conifer forest and is therefore
more difficult to find. Var. montana and var. stephensonii are listed as Critically Endangered (CR)
and var. nevadensis as Endangered (EN) on the IUCN Red List. Fire hazards are the main threat,
increased by human interference with the environment.

Xanthocyparis

Two species are recognized in this genus, one in North America and one in Asia (Farjon, 2005a;
2010a). The American species, originally described as Cupressus nootkatensis, has been juggled
about taxonomically in recent years, especially as a result of phylogenetic analyses based on
molecular data. We shall not dwell on these issues here other than to say that there is a close
relationship with Cupressus and that under a broader concept of that genus it would be included
therein (Christenhusz et al., 2011).
80 north america

Map NA-78. Xanthocyparis nootkatensis [n=71,

m=68, h=16, 1794–2006]

Xanthocyparis nootkatensis occurs along the

NW coast of North America, mainly from
Cape Fairweather in Alaska to the Olympic
Peninsula in Washington. An isolated occur-
rence has been reported from Prince William
Sound, Alaska at ca. 60° 30’ N but no herbar-
ium collection was available to put this on
our map. S of the Olympic Peninsula in Wash-
ington X. nootka­tensis leaves the coast and
occurs mainly in the Cascade Range from the
Canadian border to Josephine Co. in Oregon.
The southernmost localities are in the Klamath Mountains of extreme N California at ca. 41° 50’
N. A few isolated populations are further than 160 km (100 miles) inland in S British Columbia
and most notably at Cedar Camp on Buck Cabin Creek in Grant Co., Oregon. This species requires
high levels of precipitation and is well adapted to a cool and often cloudy climate, with conditions
at its northernmost localities in Alaska at sea level comparable to those at 1000 m and higher (to
2300 m) in the Cascade Range. Although a more or less sporadic tree in mixed conifer rain forest,
it occurs more abundantly around bogs on peaty soil and near the tree line. Xanthocyparis noot­
katensis is exceedingly long-lived, with ages verified in excess of 2000 years and estimates going
up to 3500 years for some very large, but hollow trees. Growth is very slow even in the sapling
and pole stages, with a short growing season.

Juniperus

Juniperus is a relatively large genus with 53 species (Adams, 2011 recognized 67 species) of which
11 are native in North America N of Mexico. One of these, J. communis, is the only circumpolar
conifer and is treated in the chapter Europe and the Mediterranean. In North America, species of
the genus occur from the Arctic tundra to the Californian and Texan deserts and from sea level
to alpine meadows and rocks. The genus Juniperus is as a consequence of this diversity in habi-
tats more widespread even than Pinus and occurs in every province and state of Canada and the
continental USA.

Map NA-79. Distribution of the genus Juniperus in North America and Greenland.

Juniperus is the only conifer genus with a scattered but widespread distribution in the prairie states
of the American Midwest, from North Dakota (and adjacent parts of Canada) to south-central
Texas. In Texas the genus is abundant and diverse in species. This genus has predominantly
 north america 81

adapted and diversified in response to (seasonal) aridity, reflected in this broad distribution pat-
tern. However, some are forest species, albeit not of dense, closed canopy forests, or occur in wet
coastal areas but there often on rocks or sand dunes. Arctic and alpine habitats have already been
mentioned. As a consequence of this broad ecological amplitude, the distribution of the genus
Juniperus is more evenly spread across the continent than that of other conifer genera.

Map NA-80. Juniperus horizontalis [n=151, m=142, h=19, 1779–2006]

Juniperus horizontalis is a low decumbent shrub with an essentially boreal distribution. Its extent
of occurrence reaches across the N half of the continent from Newfoundland in the E to SE Alaska
in the W. It is also present on several smaller islands in the Atlantic Ocean, the furthest out of
these is Sable Island 175 km SE of Nova Scotia. Its N limits appear to be in the Mackenzie Delta at
Cabin Creek, 69° 17’ N, second only to J. communis var. saxatilis (maps EM-31-32 on pp. 160–161).
As with other North American boreal conifers, in the E of the continent the N limits are at much
lower latitudes, but it is not well established where this line runs. The herbaria consulted (six are
in Canada) did not have collections of this species from mainland Labrador, N Quebec, N Ontario
and N Manitoba; if it occurs there it is rare. In the USA Juniperus horizontalis extends S along the
Atlantic coast as far as Virginia and in the interior an isolated locality in Stone Co., SE Missouri at
36° 48’ N is the southernmost occurrence. Although mostly a lowland species, it is not uncommon
in the foothills of the Rocky Mountains from Alberta to Wyoming where it ascends to 1160 m. Juni­
perus horizontalis does not occur W of the Rocky Mountains and in the far N follows the eastern
base and foothills of the Wrangell-Saint Elias Mountains and the northern limits of the Chugach
Mountains as far as the Kenai Peninsula, with an isolated locality on the Salcha River (Richardson
Hwy.). This indicates a preference for a continental climate in the W, in the rain shadow of the
great mountains and icefields. In the E these conditions extend to near the North Atlantic Coast,
although Newfoundland has a cool and wet climate and there this species is abundant.

Map NA-81. Juniperus scopulorum [n=124, m=120, h=19,

1804–2010]

Juniperus scopulorum has an extensive distribution

throughout the Rocky Mountains, from British Colum-
bia and Alberta in Canada to N Mexico and, sporadi-
cally, further S in parts of N Mexico. E of the Rocky
Mountains it occurs in the W parts of North Dakota,
South Dakota and Nebraska, with an isolated loca-
tion in N Texas (‘Texas Panhandle’). Here it is said to
form hybrids with J. virginiana (Adams, 2011). In British
Columbia J. scopulorum has been collected as far W as
the Bulkley River near Telkwa (R. P. Adams on 9 Nov
1976). It also occurs in the S of Vancouver Island and
82 north america

on the Gulf Islands, as well as in Puget Sound and adjacent parts of the Olympic Peninsula. These
populations were described as a distinct species J. maritima by R. P. Adams (map in Adams, 2011
p. 222). Also W of the Rocky Mountains are isolated populations in E Washington, E Oregon and
in a few mountain ranges in Nevada. The altitude at which this species is found ranges from near
sea level to 2770 m in the southern Rocky Mountains. It occupies various habitats from grassland-
artemisia steppe to mixed woodland and open conifer forest with moderate rainfall. In the Pacific
Northwest it occurs in the rain shadow of coastal mountains but in the S of its extensive range at
higher altitudes well above semi-desert conditions. Here it is a constituent of the extensive pinyon-
juniper woodlands and oak-pine forests of N Mexico.

Map NA-82. Juniperis osteosperma [n=97, m=90, h=15,

1845–2008]

Juniperis osteosperma occurs in the W of the USA from

the Rocky Mountains to the E foothills of the Sierra
Nevada and has been found in the following states:
Arizona, California, Colorado, Montana, Idaho, Nevada,
New Mexico, Utah and Wyoming. This species is most
abundant on the Colorado Plateau, a vast area of flats,
table lands, buttes and canyons and, except at the
highest altitudes, semi-desert conditions. It is also com-
mon in the Big Horn Basin of Wyoming, from where it
reaches its northernmost locality near Billings in Mon-
tana. In Arizona its southern limit is the Mogollon Rim,
a S boundary also for J. monosperma (map NA-83 on
p. 83), Pinus edulis (map NA-12 on p. 42) and Cupressus
arizonica var. glabra (map NA-77 on p. 79). In the W J. osteosperma is bounded by the Sierra Nevada
and the Mojave Desert, but there is a disjunct population in the San Jacinto Mountains, Riverside
Co., California on the far side of this desert. Very isolated populations also occur in Idaho and NE
Nevada. The altitudinal range is 460–2700 m a.s.l. Juniperus osteosperma and Pinus edulis form the
widespread pinyon-juniper woodland community, an open conifer forest type adapted to the harsh
conditions on the Colorado Plateau and inter-mountain valleys to the NE and W of this plateau, with
high summer temperatures and cold winters. At lower altitudes the juniper may form pure stands
without the pine and it is also better capable of growing from crevices in bare sandstone rock.

Juniperus osteosperma on a dry plain looking towards the canyons and mesas of the San Rafael Swell, Utah.
Photo credit Aljos Farjon.
 north america 83

Map NA-83. Juniperus monosperma [n=53,

m=50, h=12, 1874–2008]

Juniperus monosperma occurs in the

‘Four Corner States’ Arizona, New Mex-
ico, Colorado and Utah, but in the latter
state only near Zion National Park in the
SE. There is also a locality with this spe-
cies in S Nevada near Caliente, the most
western occurrence. In the E this species
is present just across the state border in the ‘Panhandle’ of Oklahoma and in the ‘Panhandle’ of
Texas, as well as in the far W of Texas. In Arizona its S limit is along the Mogollon Rim, herbarium
collections from S Arizona identified as J. monosperma should be assigned to J. arizonica (map
NA-87 on p. 84; see Adams, 2011). There is considerable overlapping in the distribution of J. mono­
sperma and J. osteosperma (map NA-82 on p. 82) and they are both constituents of the pinyon-
juniper woodland community. Juniperus osteosperma extends further N than J. monosperma and
almost all records for J. monosperma from Utah belong instead to J. osteosperma.

Map NA-84. Juniperus californica [n=74, m=67, h=19, 1859–

2009]

Juniperus californica has its main distribution in Cali-

fornia (USA) but also occurs in Baja California, Mexico
and on Cedros Island and Guadalupe Island; the latter
island is 240 km from the coast of Baja California. Fur-
thermore, two disjunct localities with small popula-
tions of J. californica occur in E Arizona. In California,
this species occurs in the dry E foothills of the Coast
Ranges and at the base of the mountains surrounding
the Mojave Desert as well as southwards in the dry
hills of San Diego Co. and into the Sierra de Juarez
across the Mexican border. Juniperus californica is one
of the most drought tolerant species of juniper and
often occurs in desert-like conditions on rocky slopes
and on gravelly flats among cacti, agaves, dwarf desert shrubs and sparse (annual) herbaceous
vegetation. The conspicuous white waxy ‘bloom’ on the otherwise reddish-brown seed cones is
an adaptation to protect the tissue from intense sunlight.

Juniperus californica on the edge of the desert in Anza-Borego Desert State Park, San Diego Co., California.
The white bloom on the seed cones protects them from the intense desert sunlight. Photo credit Aljos Farjon.
84 north america

Map NA-85. Juniperus ashei [n=67, m=64, h=16, 1872–1998]

Juniperus ashei is a juniper of the southern USA,

occurring from extreme S Missouri and N Arkansas to
N Couahuila in Mexico. The main distribution of this
species is in Texas, where it is common in the central
part of the state mainly on limestone outcroppings
and glades. Outside this central area, largely com-
prising the Edwards Plateau and upper drainage of
the Colorado River, the distribution is disjunct, with
notable populations in Garza Co., Texas, Murray Co. in Oklahoma, Sevier Co. in SW Arkansas
and more extensive populations in N Arkansas. Juniperus ashei var ovata occurs in N Coahuila,
Mexico and in adjacent Texas, where it is reported W of J. ashei var. ashei but also from within
the range of the nominate variety. Many older collections have not been separated in the herbaria
in recognition of this recently described variety and we therefore map the two together as the
species. Juniperus ashei is a multi-stemmed shrub or commonly a short stemmed tree branching
low above the base, forming extensive populations and increasingly spreading into old farm fields
that are no longer intensively cultivated.

Map NA-86. Juniperus pinchotii [n=64, m=60, h=16, 1847–

1999]

Juniperus pinchotii occurs in Texas and adjacent

parts of Oklahoma (Roger Mills Co.), New Mexico
(Chaves Co., Eddie Co. and Lincoln Co.) and N Coa-
huila, Mexico. It has also been collected in mountains
near Galeana in Nuevo León, a disjunct population
ca. 400 km S of the main distribution. It is common to
abundant across much of central and W Texas on hills
and in ravines and canyons, often but not exclusively
on limestone and gypsum. Juniperus pinchotii invades
grassland areas that are no longer grazed intensively
and is considered a weed by farmers. In the W and S
this juniper is more often a constituent of pinyon-juni-
per woodland and in Nuevo León it has been found in
open pine-oak woodland at 1900–2150 m altitude.

Map NA-87. Juniperus arizonica [n=31, m=31, h=12, 1800–1988]

Juniperus arizonica is closely related to J. monosperma

(map NA-83 on p. 83) and J. pinchotii (map NA-86
on p. 84) as well as to the Mexican junipers J. angos­
turana (map MCW-45 on p. 117) and J. coahuilensis
(map MCW-46 on p. 117). All have (predominantly) a
single seed in the seed cones. Juniperus arizonica was
described and named by R. P. Adams in 2006 but many
herbarium specimens that should be assigned to this
species are still labelled as J. erythrocarpa or J. pinchotii
and care must be taken in cases where these are made available in databases on the Internet, e.g.
via GBIF. Juniperus arizonica occurs in central and S Arizona and in SW New Mexico and accord-
ing to Adams (2011) also in NE Sonora, Mexico. Herbarium collections in and from Mexico are in
need of taxonomic revision before they can be used with confidence for mapping, hence the single
dot on our map.
 north america 85

Map NA-88. Juniperus occidentalis var. occiden­ Juniperus occidentalis var. australis below Kaiser Pass
talis (black) [n=69, m=66, h=16, 1825–2006]; in the Sierra Nevada, California. Photo credit Aljos
J. occidentalis var. australis (red) [n=39, m=39, h=15, Farjon.
1845–2003].

Juniperus occidentalis occurs in the western states of Washington, Oregon, Idaho, California and
Nevada. It is widespread in Oregon and California but its distribution is more scattered in the
other states, particularly Idaho and Nevada. In Oregon it reaches the coast in a few places, but
otherwise it is an inland species, becoming most abundant E of the Cascade Range, the Siskiyou
Mountains and in the Sierra Nevada. Two varieties are recognized which appear to be separated
geographically, with var. australis in the Sierra Nevada, the Panamint Mountains and the San Ber-
nardino Mountains of California and a few localities in W and S Nevada. This variety was raised
by R. P. Adams to species rank and named J. grandis (‘Grand juniper’) and is indeed remarkable
for the great size (and age) some of these high altitude (1980–3100 m) trees have attained. They
occur often in places where bare granite rock sculpted by glaciers dominates the landscape and
junipers and pines grow from crevices or in ancient rock slides, slowly but out of reach of forest
fires. In contrast, J. occidentalis var. occidentalis appears to mostly occupy an ecotone between
sagebrush (Seriphidium tridentatum) steppe and Pinus ponderosa forest at much lower altitude
(200–1200 m). Fires will prevent it from growing very large and in the absence of fire pines and
other conifers will replace it. Unlike the situation with var. australis, which regenetates sporadi-
cally and slowly, var. occidentalis is now aggressively spreading on to range lands largely due to
changes in the vegetation caused by overgrazing in the past.
86 north america

Map NA-89. Juniperus virginiana var. virginiana (black) [n=134, m=125, h=18, 17??–2010]; J. virginiana var.
silicicola (red) [n=25, m=24, h=7, 1832–2006]

Juniperus virginiana is one of the most widespread species of juniper in North America, but is
limited to the E half of the continent and largely to the USA. In Canada it only occurs in the
southernmost parts of Ontario and Quebec. Two varieties are recognized; var. virginiana and var.
silicicola. The latter is a coastal tree in the SE of the USA, occurring from North Carolina S to the
coasts of central Florida and E to the Mississippi River Delta around Lake Pontchartrain. This
coastal variety occupies dunes and river sand banks, usually near their mouths and is rarely found
more than 10 km from the ocean. It is geographically well separated from var. virginiana, although
the nominal variety reaches the coast further N as well as in Texas. Juniperus virginiana var. vir­
giniana occurs far inland and has spread to the prairie states from South Dakota to Texas. Its N
limits are determined by the duration and severity of winter. In the W it meets J. scopulorum e.g.
in South Dakota (map NA-81 on p. 81) and apparently forms hybrid swarms; the same can occur
with J. horizontalis (map NA-80 on p. 81) along the N margin of its range. Juniperus virginiana is
one of the species of juniper that can become aggressively weedy, invading abandoned fields and
roadsides. This ‘weediness’ is of course a subjective term; since both farm fields and roadsides are
not natural habitats and J. virginiana is a native species, an ecologist would see this as a pioneer
conifer reclaiming lost habitat in an early phase of natural reforestation.

Calocedrus

The genus Calocedrus has a disjunct distribution with one species in W North America and three
species in E Asia. A map of the distribution of the genus is presented in the chapter Mainland
Asia and Japan.

Map NA-90. Calocedrus decurrens [n=108, m=103, h=21, 1800–2006]

Calocedrus decurrens occurs in Oregon and California and in a small

W corner of Nevada, as well as in the N part of Baja California,
Mexico. In Oregon and N California it is spreading from the Coast
Ranges across the Cascade and Siskiyou Mountains well into the
interior, but eventually becomes rare and then disappears further
E because of too little annual rainfall. For the same reason we find
it further S only in the Sierra Nevada and in the highest mountains
of S California; in Mexico it becomes rare and is known from as
far S as the Sierra San Perdro Martír, the southernmost location of
a mixed conifer forest which, albeit less diverse, has conifer spe-
cies also found in Oregon and California. This mountain system is a
 north america 87

high ‘island’ (summit 3096 m) of coniferous forest in a ‘sea’ of coastal chaparral and desert scrub,
falling steeply off to the E on to below sea level salt flats. While on this and other mountains in
the S part of its range it occurs at high altitudes (to 2960 m), in the N it can be found as low as
50 m though commonly from 500–1500 m, where it can be a tall tree with a maximum of 70 m.
Although associated with many species of conifer, C. decurrens has a preference for light, open
woodland settings and often occurs on sunny, rocky slopes and ridges, sometimes stunted to
dwarf trees just a few metres tall.

Taxaceae

The family Taxaceae is represented in North America by two genera, Taxus with three species
and Torreya with two species. Compared to temperate mainland Asia, which has four genera
(plus Cephalotaxus, here treated in a separate family) and many more species, North America is
‘impoverished’ but not as severely as Europe, which has only a single species of Taxus. The latter
genus has a wide distribution in the Northern Hemisphere, reaching into the tropics and S of the
Equator in Malesia. Torreya is much more limited and scattered; in North America in California
and Florida, and in Asia it is restricted to China and Japan. Species of Taxaceae are commonly
understorey shrubs or small trees, but Taxus can under certain edaphic conditions colonize open
grass or scrub vegetation.

Map NA-91. Taxus brevifolia [n=96, m=96,

h=15, 1860–2006]

Taxus brevifolia occurs in the W of North

America from extreme S Alaska and
the Queen Charlotte Islands of British
Columbia along the coastal islands and
mountains to N California. There is a
second area more inland in the Selkirk
Mountains, British Columbia and in NW
Montana, N Idaho, NE Washington and
NE Oregon. In California, the distribution
continues S through the Sierra Nevada
where it reaches its southernmost locali-
ties in Calaveras Co. This distribution
pattern is similar to that of several coni-
fer species in the NW of the continent,
i.e. Tsuga heterophylla (map NA-50 on p. 62), Abies grandis (map NA-57 on p. 66), Thuja plicata
(map NA-69 on p. 74) and to a lesser extent T. mertensiana (map NA-51 on p. 63). This pattern is
strongly determined by climatic factors, in particular high rainfall and cool summers. The ever-
green coniferous temperate rain forest of the coastal region extends inland to those western areas
of the Rocky Mountains which receive similar high rainfall coming in from the Pacific Ocean. It
is obvious that these species will be most abundant on the wetter and cooler NW and W sides
of mountains, but the detail of our maps is not sufficient to show this. What is clear is that the
region E of the Cascade Range for a distance of 150–200 km is too dry for Taxus brevifolia, which
in nearly all situations is an understorey shrub or tree in conifer forest, but can occur on rocky
outcrops at sufficiently high altitude or close to the coast, e.g. in the Queen Charlotte Islands. As
with other conifers, the gap along the British Columbia Coast is not real but an artefact of under-
collecting in that region.
88 north america

Map NA-92. Taxus canadensis [n=130, m=129, h=10, 1871–2008]

Taxus canadensis is the eastern vicariant of T. brevifolia, occurring from Newfoundland in the E
to Manitoba in the W. The shortest distance between the two species is ca. 1300 km. The north-
ernmost locality of T. canadensis is at York Factory on the SE coast of Hudson Bay (57° N 92° 18’
W); to the S it reaches the Appalachian Mountains in North Carolina and Tennessee (36° 24’ N).
The population at Hudson Bay and another on Hecla Island in Lake Winnipeg (at 96° 43’ W
the furthest locality W with a herbarium record) are outliers; the main distribution in the NW
surrounds Lake Superior. To the S the Mississippi Valley in Wisconsin and Iowa forms the W limit
of T. canadensis. This species occurs in both coniferous and angiosperm forest as well as in mixed
forests. Its tolerance of low light levels enables it to grow under canopy of both Fagus grandifolia
and Tsuga canadensis. In other situations this layering and spreading shrub can grow outside for-
est cover in bogs or moist rocky ravines, where it avoids competition.

Map NA-93. Taxus floridana [n=13, m=12, h=5, 1833–1989]

Unlike the two other North American species, Taxus

floridana has an extremely limited distribution in
NE Florida along the Appalachicola River and a few
tributary streams in Gadsden and Liberty Counties.
It occurs in moist ravines and on limestone bluffs in
forest or woodland dominated by Fagus grandifolia,
Magnolia grandiflora and Quercus laurifolia, often
with several other trees present, among which occa-
sionally another narrowly endemic conifer, Torreya
taxifolia (map NA-95 on p. 89). On bluffs it may occur
in thickets of Kalmia latifolia. These woodland species do not occur in natural vegetation sur-
rounding these ravines (mostly dominated by Pinus elliottii and P. palustris) and strongly indicate
that these ravines shelter a relict forest with past links to the Appalachian Mountains and thus
with Taxus canadensis, and that T. floridana could have evolved from southern populations of
that species. However, in one phylogenetic analysis using the ITS region of nuclear ribosomal
DNA (Li et al., 2001) T. floridana was most closely related to T. globosa of Mexico and Central
America. The habitat of T. floridana is under stress from drought caused by agricultural practices
on the surrounding land, increasing the risk of fires as well as pests. This species is listed as Criti-
cally Endangered (CR) on the IUCN Red List.
 north america 89

Map NA-94. Torreya californica [n=58,

m=58, h=13, 1851–2006]

Torreya californica occurs in two sepa-

rate regions, divided by the Central
Valley of California. The most western
region comprises the Coast Ranges N of
San Francisco as far as Trinity Co. and
the Redwood forests along the coast
from Mendocino Co. S to Santa Cruz
Co. The region E of the Central Valley
is on the W slopes of the Sierra Nevada
along its entire length. An isolated most
northern occurrence is at Lassen Buttes
in Plumas Co. Torreya californica is thus
endemic to the state of California. It is
everywhere an uncommon or rare under-
storey tree usually growing in moist and
shady places in hilly to mountainous
terrain dominated by tall conifers, espe-
cially Sequoia sempervirens and Pseudot­
suga menziesii. In the the Sierra Nevada
it is associated with angiosperms, mainly
Acer spp., Cornus nuttallii, Platanus occidentalis, and Alnus rubra and here it strictly adheres to
stream banks in steep canyons. Its altitudinal range is from near sea level on the coast to 2500 m
in the Sierras, but most commonly it occurs between 200 m and 1200 m. Exploitation in the past
of this slow growing tree has caused substantial decline and the species is listed as Vulnerable
(VU) on the IUCN Red List.

Map NA-95. Torreya taxifolia [n=19, m=17, h=5, 1843–1980]

Torreya taxifolia is rare and extremely localized,

occurring in the valley of the Appalachicola River in
extreme SE Georgia and NE Florida, for a distance
of 64 km. Its distribution is similar to that of Taxus
floridana (map NA-93 on p. 88) but extending a short
distance futher upstream across the Florida/Georgia
border. A disjunct occurrence 11 km W of the river in
Jackson Co., Florida is not shown on the map as no
herbarium collection from this locality was available.
Torreya taxifolia grows mostly in the shade of wooded
ravines and steep, N-facing slopes under canopy of
Fagus grandifolia, Liriodendron tulipifera, Magnolia grandiflora, Acer barbatum, Liquidambar
styraciflua, Quercus alba, and occasionally pines (Pinus taeda, P. glabra). Sometimes it is found
together with Taxus floridana and both are undoubtedly relicts of a wider distribution in the geo-
logical past. Both species may have occurred in the southern Appalachian Mountains before the
last (Wisconsin) glacial maximum but failed to migrate back when this cold phase ended some
10,000 years ago. Torreya taxifolia is attacked by an introduced pathogenic ascomycete (fungus)
and most of the fewer than 1000 remaining live individuals are now resprouts from stumps. This
species is listed as Critically Endangered (CR) on the IUCN Red List.
MEXICO, CENTRAL AMERICA AND THE WEST INDIES

Geography

This region includes Mexico, Guatemala, Belize, Honduras, El Salvador, Nicaragua, Costa Rica
and Panama on the North American mainland and Cuba, Jamaica, Hispaniola, Puerto Rico, the
Bahamas and the Lesser Antilles as islands in the Caribbean Sea. Included are also two oceanic
islands: Guadalupe Island in the Pacific Ocean and Bermuda in the Atlantic Ocean. Its geographi-
cal boundaries are on the whole natural except for the artificial one between Mexico and the USA.
It is very diverse geographically, with a subcontinent (Mexico), an isthmus (Central America) and
large and small islands in a tropical sea (the Caribbean) and in this chapter we will treat these
three parts of the region separately.
The total land area of this region is approximately 2.75 million km² of which 2.52 million km²
is on the mainland and only 0.235 million km² comprises the islands. Much of the region lies
in the tropics, but Mexico extends N to ca. 32° N, the Bahamas to 27° N and Bermuda is at
32° 19’ N; the Panama/Colombia border is the region’s farthest point S at ca. 8° N. The mainland
is mountainous across the entire length from the USA border to the border between Panama and
Colombia, but with a few (near) gaps of which the Isthmus of Tehuantepec in S Mexico is the
most significant. The Caribbean islands are also mountainous, with the exception of the very flat
and low Bahamas and extensive lowlands in Cuba and the Dominican Republic (the eastern half
of Hispaniola). In Mexico the main mountain ranges are the Sierra Madre Occidental and Sierra
Madre Oriental, the Transverse Volcanic Belt and the Sierra Madre del Sur. In Central America
the highlands are more or less continuous in Guatemala, El Salvador and Honduras, but break up
in sections in Nicaragua, Costa Rica and Panama.
The climate in the West Indies is tropical, with prevailing NE trade winds and oceanic influ-
ence moderating temperatures but also bringing in hurricanes generated over the tropical Atlan-
tic. Mountains influence temperature and above all precipitation, with the windward slopes
much wetter than the leeward slopes. On the mainland, climate is much more diverse, from
desert conditions in parts of N Mexico to cold alpine on the highest summits. Warm temperate
and tropical montane rainforest occurs along the Gulf of Mexico and in parts of S Mexico and
Central America, where the lowlands are hot tropical. In the West Indies as well as in S Mexico
and Central America there are two seasons, a rainy season from May through October and a dry
season during the remaining months, corresponding with winter at more northerly latitudes. In
N Mexico rain is sporadic or erratic and occurs as summer thunderstorms, except in Baja Califor-
nia, which has three climate zones: tropical at the southern end, desert conditions for the greater
length, and winter rainfall in the northern parts including the Pacific islands.

Conifers in Mexico, Central America and the West Indies

Families Genera Species Species + infra Countries with Area of occupancy Endemic taxa
conifers in km²
4/8 11/70 99/615 132/794 23/34 55,525/2,751,100 100
Numbers after ‘/’ are global or continent totals.
 mexico, central america and the west indies 91

This region is relatively diverse in conifers, with half of the world’s eight families, 11 genera (16%),
99 species (16%) and 132 species + subspecies and varieties (16%). Of these 100 (75%) are endemic
to the region. Of the 32 non-endemics, 29 also occur in the USA and only three are shared with
South America. The border with the USA being not only artificial but also very long, it is inevi-
table that some taxa have their main distribution in the USA and just happen to occur across the
border into Mexico. These are counted here, but are discussed and mapped in the chapter North
America. All eight countries on the mainland have native conifers and all the larger islands and
archipelagos in the West Indies have conifers; the 11 countries (or dependencies) lacking conifers
are all very small islands or groups of islands. The more notable (near) absences of conifer spe-
cies in the region are in the Chihuahuan and Sonoran Deserts, in the lowlands of Tamaulipas and
the Yucatán Peninsula (all Mexico), in the southern half of Nicaragua and in the lowland areas of
Cuba and Hispaniola. The lowlands of Nicaragua not only form a gap in conifer distribution overall,
here is the southern limit for Northern Hemisphere conifers in the Americas with only Cupres-
sus and Pinus actually reaching it. The genus Pinus (Pinaceae) is the most diverse group; with 47
species, total of 61 species + infraspecific taxa this region is the world’s centre of diversity of the
genus (Farjon & Styles, 1997). Another diverse genus is Juniperus (Cupressaceae) with 21 species,
total of 31 species + varieties. There are 13 species of Podocarpus (Podocarpaceae) in this region.
These three genera are the only ones also present on the Caribbean Islands, the remainder of coni-
fer diversity occurs on the mainland only. These are Abies (Pinaceae) (7, 12), Cupressus (Cupres-
saceae) (3, 7), Picea (Pinaceae) (3, 3), Calocedrus (Cupressaceae) (1), Prumnopitys (Podocarpaceae)
(1), Pseudotsuga (Pinaceae) (1, 1), Taxodium (Cupressaceae) (1) and Taxus (Taxaceae) (1).

Map MCW-1. Distribution of conifer species in Mexico, Central America and the West Indies.

The taxonomic diversity of conifers both at generic and species levels is highest in the Sierra
Madre Oriental and in Oaxaca and Chiapas. With the exception of an area in Durango, highest
numbers of genera (7) and species (>18) per 0.5 degree cell are found in the mountains facing the
Gulf of Mexico, where precipitation averages are highest. Central America and particularly the
West Indies have lower diversity values. The desert margins of N Mexico (Chihuahuan & Sonoran
Deserts) are also poor in conifer diversity. At the species level, diversity is somewhat more dis-
persed yet shows the same general pattern as for the genera. The high diversity in the genus Pinus
accounts for most of the high diversity cells, here and there supplemented by species of Juniperus
and to a lesser extent Abies.
The rarity calculation illustrated in Map MCW-4 shows narrow endemism to be more wide-
spread in the region and includes parts of the West Indies and Guadalupe Island in the Pacific
Ocean. The island effect on endemism is apparent here in higher scores than in the diversity
92 mexico, central america and the west indies

maps, but the highest score of one cell in Hispaniola is merely due to a localized variety: Junipe-
rus gracilior var. urbaniana (map MCW-66 on p. 131). The E of Mexico near the Gulf which has
the highest diversity also has most of the local endemism, including the highest scoring cells
in SE Coahuila and Nuevo León. Some high mountains are in effect ecological islands causing
endemism and we find examples in several parts of Mexico, including the three cells with the
highest score.

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�‒�
�
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Map MCW-2. Map of Mexico, Central America and the West Indies showing numbers of conifer genera
per 0.5 degree cell.

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�‒�
��‒��
��‒��
��‒��

Map MCW-3. Map of Mexico, Central America and the West Indies showing numbers of conifer species +
infraspecific taxa per 0.5 degree cell.

Map-MCW-4. Rarity values for 0.25 degree cells. Cell score is the total of the rarity calculations for each
species in the cell. The rarity calculation is 1 divided by the total of cells occupied by a species. Thus, a spe-
cies that occurs in one cell scores 1 while a species that occurs in 10 cells scores only 0.1.
 mexico, central america and the west indies 93

Mexico

Geography

Mexico is the largest country in the region with a land area of 1.97 million km². It extends from
32° 32’ N to 14° 32’ N and its border with the USA is ca. 3130 km long. In the NW Baja California
forms a peninsula with a length of 1247 km, separated from mainland Mexico by the still widening
(through sea-floor spreading) Gulf of California. In tropical latitudes the large Yucatán Peninsula
extends into the Caribbean Sea and, unlike Baja, is a low-lying plain. Major mountain ranges
make up much of the rest of Mexico. In the N these are the Sierra Madre Occidental and Sierra
Madre Oriental; between these more or less NNW-SSE oriented systems stretches a vast, elevated
basin, the Altiplano Mexicano. To the S of these the Transverse Volcanic Belt (Eje Volcánico
Transversal) reaches from coast to coast, with several volcanoes higher than 5000 m, culminating
in Pico de Orizaba (Citlaltépetl) near its eastern end, the highest mountain of the entire region at
5636 m. Further S the Sierra Madre del Sur rises along the Pacific Coast and beyond the Isthmus
of Tehuantepec the Sierra Madre de Chiapas connects with the Guatemalan Highlands. With
more than half of the country’s area above 1000 m, high altitude is a predominant feature relevant
to conifers, yet besides the Yucatán other extensive lowlands occur especially along the Gulf of
Mexico. Narrower strips of lowland are situated along the Pacific coast and the mainland side of
the Gulf of Mexico. The geology of Mexico is complex and cannot be set out here in any detail;
important features are the Eocene-early Oligocene uplift of the Sierra Madre Occidental, the vol-
canic origin of the Sierra Oriental and, of younger age, the Transverse Volcanic Belt, and the fact
that much of the uplift as well as the origin of major volcanoes in areas of high conifer diversity
date from relatively recent (Miocene through Pleistocene) geological periods. The aridification of
the N of Mexico is partly a result of these orogenetic events and has had a major impact on the
development of the flora, including conifers. The Chihuahuan and Sonoran Deserts, with a few
local exceptions, are not extreme deserts and are relatively well vegetated, especially with suc-
culent cacti, yuccas and agaves. Along both flanks of the Sierra Occidental and along the W side
of the Sierra Madre Oriental the deserts give way first to dry open (deciduous) woodland and at
higher altitudes to mixed oak-pine forest and ultimately pure conifer forest. On the E side of the
Sierra Madre Oriental and on the N side of the Sierra Madre del Sur cultivated lowland merges
upward to rain or cloud forest dominated by angiosperms in which conifers occur more sporadi-
cally. Conifers dominate most of the montane to subalpine forests in the Transverse Volcanic Belt.
At lower elevation both here and on the S flank of the Sierra Madre del Sur tropical deciduous
woodland, sometimes rich in large cacti, dominates. Unlike Central America and some islands
in the West Indies, Mexico does not have extensive low lying coastal pine savannas. Secondary
tropical lowland rainforest, seasonally dry tropical forest and thorn scrub occupy large parts of
the Yucatán Peninsula, with only two small marginal populations of Pinus caribaea.

Conifers in Mexico

Families Genera Species Species + infra Countries with Area of occupancy Endemic taxa
conifers in km²
4/8 11/70 80/615 108/794 1/1 39,825/1,972,545 61
Numbers after ‘/’ are global or country totals.

Mexico is the country with by far the greatest diversity of conifers in the region, with all 11 genera,
80 species and 108 species + infraspecific taxa, of which 61 are endemic. More than half of these
species + infraspecific taxa, 44 species and 56 species + infraspecific taxa, belong to the genus
Pinus, for which Mexico is the world’s centre of diversity (Farjon, 1996). Juniperus accounts for
94 mexico, central america and the west indies

17 species and 24 species + varieties, Abies (7, 12), Cupressus (3, 7), Picea (3, 3), Podocarpus (3),
Calocedrus (1), Pseudotsuga (1, 1), Taxodium (1) and Taxus (1). In Mexico, the distribution of coni-
fer species is concentrated in the mountains. Pinus muricata is the only species that is restricted
to low altitudes along the Pacific coast of northern Baja (and in California, USA). The lowlands
of Mexico, with few exceptions (e.g. an occurrence of Pinus caribaea var. hondurensis at 120 m
in Quintana Roo, Yucatán Peninsula) are devoid of naturally occurring conifers. In the Chihua-
huan and Sonoran Deserts conifers occur only on isolated mountains or along streams (Taxodium
mucronatum) mostly with permanent (ground) water. While conifers are nearly ubiquitous in the
mountains, pure conifer forests are limited to the higher altitudes and virtually absent in tropical
deciduous forest. The vast majority of forests are mixed, mostly oak-pine (the genus Quercus is,
like Pinus, richest in species in Mexico) but more diverse in the rain and cloud forests around the
Gulf of Mexico and in Oaxaca. Map MCW-5 shows concentrations that are the result of species
diversity in these areas; to a large extent this is accounted for by the genus Pinus, with often as
many as 10 species on a single mountain. Mountain areas dominated by deciduous tropical for-
est, in which a prolonged dry season is evident, also have few if any conifers. These forests are
particularly extensive along the Pacific coast of Mexico and occur at low to middle altitudes; the
highest parts of the mountains usually have forests with pines or sometimes other conifers.

Map MCW-5. The distribution of conifer species in Mexico.

Pinus

The distribution of the genus Pinus (Pinaceae) is very similar to that of all conifer species in
Mexico. This is not surprising as the taxonomic diversity of the genus in Mexico reflects adap-
tations to virtually all types of habitat in which conifers occur in that country. Exceptions are
semi-arid grasslands in the NE fringing the Chihuahuan Desert where some species of Juniperus
are usually the only conifers. However, these grasslands merge at somewhat higher altitudes with
pinyon-juniper woodland, where the genus Pinus meets its limits to aridity. At the other end of
the precipitation scale, the genus Pinus is rare or absent in the wettest montane rainforests and
in lowland tropical rainforest. Competition with angiosperm trees limits pines to scattered small
 mexico, central america and the west indies 95

areas of exposed ridges and summits with thin or skeletal soils. A total of 44 species of Pinus is
currently recognized here, and while this genus is comparatively well studied, new species are
being described, often as minor segregates from already known species. A number of these are
here recognized as subspecies and varieties, taking the total of taxa in Pinus in Mexico to 56
(Farjon & Styles, 1997; Farjon, 2010a).

Map MCW-6. The distribution of species in the genus Pinus in Mexico.

The uncertainty about the number of species is partly due to the existence of widespread poly-
morphic species or species alliances and complexes that pose challenges to species delimitation.
A complex geological and climatological past has caused diversification, extinction and migration
so that at present the taxonomic diversity is distributed unevenly. Some species have enormous
ranges right across the country and beyond, others are extremely limited and either recent neoen-
demics or possibly relict populations of a once wider distribution. As observed above, taxonomic
diversity can be localized, with many species occurring in one location, e.g. a mountain, even if
altitudinal zonation often separates species. On a wider scale taxonomic diversity of Mexican
pines is unevenly distributed, with several centres (see map MCW-6).
Some of these centres of high diversity are situated within the two regions of speciation out-
lined by Farjon (1996) but others appear to occur elsewhere. Overall diversity does not necessarily
equate with centres of speciation because relict and presumably ancient species are included in
the calculations. A phylogeny of the genus can indicate to some extent which species represent
more recent lineages if lineages can be dated approximately. While many phylogenetic analyses
have been published (e.g. Gernandt et al., 2005), some testing hypotheses based on the fossil
record (Eckert & Hall, 2006), ‘molecular clock dating’ with calibration by the fossil record has
only begun recently (Gernandt et al., 2008) and these results do not yet enable us to separate the
recent from the more ancient species of Pinus in Mexico. General diversity has to indicate centres
of speciation including the ‘noise’ from more ancient lineages.
A substantial number of species in Pinus have their main distribution N of the Mexico/USA
border and extend into Mexico; these are mapped and discussed in the chapter North America.
This includes Pinus strobus var. chiapensis, which occurs in S Mexico and Guatemala but are
96 mexico, central america and the west indies

mapped with P. strobus var. strobus to show the remarkable, but not improbable, disjunction
in this species. Also in the south, several species with wide occurrences in Mexico extend into
Central America; these are mapped and discussed here. The exception is Pinus caribaea var. hon-
durensis, which although mostly occurring in Central America, are treated with the other two
varieties in the section on the West Indies within this chapter. No pine species in Central Amer-
ica are entirely absent from Mexico. In the following account the species of pines are grouped
according to their classification in two subgenera: Pinus and Strobus.

Subgenus Pinus

Map MCW-7. Pinus oocarpa [n=167,

m=164, h=41, 1838–1994]

Pinus oocarpa has the widest distri-

bution of all Mexican pines, extend-
ing over a linear distance of ca. 3000
km from the border area between
Sonora and Sinaloa in NW Mexico
to the Cordillera Isabella in Nicara-
gua. It is less common in the Sierra
Madre Occidental and there are gaps
in parts of Michoacán and Guerrero
and also in Puebla and W Oaxaca.
Extensive pine forests dominated by
P. oocarpa are mostly concentrated in Guatemala, Honduras and NW Nicaragua. It has also a wide
altitudinal range, from 200 m to 2700 m and occurs under widely different climatic conditions,
with e.g. annual precipitation between 700 mm and 3000 mm (Farjon & Styles, 1997). Its optimum
lies around 1000–1500 m a.s.l. with abundant rainfall, but with a long dry season from October to
June in much of its range. With its semi-serotinous cones P. oocarpa is adapted to fire and, with
other species of pine and/or oaks (Quercus) forms open forests or woodlands. Where fires are
frequent, grasses or sometimes ferns (Pteridium aquilinum) form the main undergrowth; if less
frequent, shrubs will dominate and can slow down regeneration when becoming too dense.

Map MCW-8. Pinus luzmariae (black)

[n=167, m=164, h=41, 1838–1994];
P. prae­termissa (red) [n=26, m=26, h=16,
1892–1991]

Pinus luzmariae and P. praetermissa

are closely related to P. oocarpa and
were formerly included, the first
species as P. oocarpa var. trifoliata.
The distribution of P. luzmariae is
probably imperfectly known and
although it is much less common,
seems to range almost as widely as
P. oocarpa. Its ecology, as far as is
known, also matches that of P. oocarpa although the altitude range as recorded from herbarium
specimens is narrower, from 820 m to 2500 m a.s.l. Pinus praetermissa has a better defined range
and is most common in W Central Mexico, especially Jalisco but extending into some neighbour-
ing states. The altitude ranges from 900 to 1900 m, much more limited and also lower than either
P. luzmariae or P. oocarpa. This is one of a few pines that can thrive in the deciduous tropical
 mexico, central america and the west indies 97

forest at middle altitudes which is dominated by angiosperms and mostly bare of foliage during
the long dry season. The pines are among the few evergreens in this ecosystem.

Map MCW-9. Pinus pseudostrobus var. pseudostrobus (black) [n=147,

m=137, h=40, 1838–1994] and Pinus pseudostrobus var. apulcensis (red)
[n=115, m=111, h=29, 18??–1994] Pinus pseudostrobus var. apul-
censis in Oaxaca, Mexico. Photo
credit Aljos Farjon.

Pinus pseudostrobus is nearly as widely distributed as P. oocarpa, ranging over a distance of 2500
km from N Mexico to Honduras and unlike that species extends far N along the Sierra Madre Ori-
ental. Pinus pseudostrobus, here interpreted in a wider sense and with two major varieties based on
seed cone morphology, is extremely variable and probably constitutes a ‘species complex’ (Stead,
1983) involving ongoing speciation. The two varieties mapped here have only partly overlapping
ranges, but especially in the southern parts can occur sympatrically, with many intermediate
forms (Farjon & Styles, 1997). Pinus pseudostrobus var. apulcensis (syn. P. oaxacana) overlaps in S
Mexico and Central America with var. pseudostrobus but in Central and N Mexico it is limited to
the eastern part of the Transverse Volcanic Belt and the Sierra Madre Oriental. Pinus pseudostro-
bus var. pseudostrobus also occurs in the central and W parts of the TVB and is sporadically pres-
ent in the Sierra Madre Occidental as well as in the Sierra Madre Oriental. Both taxa are absent
in the Isthmus of Tehuantepec, where some mountains rise to 1200–1400 m which is at the lower
end of the altitude range of the species (850–3250 m). Pinus pseudostrobus is abundant in many
mixed pine forests, pine-oak forests or at its highest altitudes it can be associated with Abies.

Map MCW-10. Pinus hart­wegii [n=149,

m=135, h=33, 18??–1994]

Pinus hartwegii grows at high alti-

tudes (2300–4300 m) across its wide
range in Mexico and Central America,
commonly forming the tree line on
the higher summits. It is particularly
common and abundant, often form-
ing pure stands, across the Transverse
Volcanic Belt of central Mexico and
in the Guatemalan Highlands. On the
highest mountains (max. 2850 m) in
98 mexico, central america and the west indies

Honduras it forms small conifer forests on the summits, often mixed with Cupressus lusitanica,
Abies guatemalensis or Taxus globosa. Below the pure tree line stands it occurs with other pines,
e.g. P. montezumae and P. ayacahuite, and Abies spp. or Cupressus lusitanica. Above 3500 m snow in
winter can last several months and on the highest volcanoes that have small summit glaciers some
straggling and dwarfed individuals of P. hartwegii grow not far below the permanent snow line.

Pine forest of pure Pinus hartwegii at around 4000 m altitude on Mexico’s highest mountain, Pico de Orizaba
(Citlaltépetl) in Veracruz (5636 m). Photo credit Aljos Farjon.

Map MCW-11. Pinus maximinoi [n=145, m=134, h=42, 1841–1994]

Pinus maximinoi is widespread from the Transverse Volcanic Belt to NW Nicaragua. Its great-
est abundance is in Guatemala and Honduras. A distant outlying population was sampled by
M. Martínez in Sinaloa (red circle). Its altitudinal range is great: 450–2800 m, with an optimum
at 900–1800 m a.s.l. It is often a constituent of pine or pine-oak forests and occurs with many
other species of Pinus, both in primary forest and in secondary forest. Pinus maximinoi has a wide
ecological amplitude from relatively dry pine-oak forest in Central Mexico to wet subtropical
 mexico, central america and the west indies 99

montane forest and cooler high altitude cloud forest in Central America. With such wide tol-
erance to ecological conditions the absence of this species in large areas in Central Mexico is
difficult to explain. The northern limit, except for the outlier in Sinaloa, is the N flank of the
Transverse Volcanic Belt and is probably a winter temperature limit. Precipitation levels in Cen-
tral America are reported to lie between 900 and 2500 mm per annum, so perhaps the long dry
season that occurs in some areas in Central Mexico could account for its absence.

Map MCW-12. Pinus tecunumanii [n=153, m=143, h=31,

1842–1996]

Pinus tecunumanii is a tropical forest pine com-

mon in Chiapas, Mexico and in Guatemala, Belize,
El Salvador, Honduras and NW Nicaragua. A dis-
junct population occurs in Oaxaca, Mexico, W of
the Isthmus of Tehuantepec. Its altitudinal range is
considerable: 300–2900 m, but it occurs in locations
with at least 1000 mm precipitation per annum and
often 2–3 times more than this. Yet a long dry sea-
son from November to May applies, during which
fires are common, but usually only destroying the undergrowth of grasses, ferns (Pteridium) or
shrubs. Pinus tecunumanii often grows associated with other pines, e.g. P. oocarpa, P. maximinoi,
and P. pseudostrobus, and at the more mesic sites P. ayacahuite and P. strobus var. chiapensis. Oaks
(Quercus) are the most common angiosperm trees, but a more diverse assembly of broad-leaved
trees is found on the wettest mountain slopes facing the Caribbean Sea, from where most of the
rain originates. Pinus tecunumanii is an important tree in tropical forestry and attempts have been
made to introduce it in many countries, so far with limited commercial success. Logging pressures
have caused this species to be listed as Vulnerable (VU) on the IUCN Red List.

Map MCW-13. Pinus douglasiana [n=95, m=94, h=24,

1897–1994]

Pinus douglasiana occurs in the mountains of

Mexico W of the Continental Divide, from south-
ern Sinaloa in the north to Oaxaca in the south. In
Morelos are the only occurrences of this pine that
straddle this great divide, separating drainage into
the Pacific from that into the Atlantic. Altitude
ranges between 1100 m and 2700 m and precipita-
tion averages at 1000 mm in its optimal altitude
range of 1400–2500 m. This species occurs in mixed
conifer forest, in pine forest (where it associates with other species of Pinus) or in pine-oak forest.

Map MCW-14. Pinus jaliscana [n=27, m=24, h=15, 1900–1994]

100 mexico, central america and the west indies

Pinus jaliscana is endemic to the state of Jalisco, where it occurs in the NW part of the Sierra
Madre del Sur, on the Pacific slope, mainly in the Sierra de Cuale (Sierra el Tuito) and S of Villa
Purificación. This is a species of moderate altitude (800–1650 m) in mountains with a prolonged
dry season from December to May. Low montane subtropical forest dominated by oaks (Quercus
spp.) or pine-oak forest with the more common species P. maximinoi and P. oocarpa on acidic soil
are its most widespread habitats. At the highest altitudes it can be joined by P. douglasiana.

Map MCW-15. Pinus patula [n=124, m=115, h=46, 1828–1994] Pinus patula in the Sierra de Juarez,
Oaxaca, Mexico. Photo credit Aljos
Farjon.

This species is distributed in E and S Mexico and is endemic to that country. With its long, slender
and drooping needles, this is a pine of moist climate, which partly explains its general distribu-
tion. Annual precipitation is from 1000 mm to 2200 mm at an altitude range of 1500 m to 3000 m
in warm to cool temperate conditions. The dry season is less severe in most of its range and nearly
absent at high altitude in cloud-forest, when fog replaces rain during this time. Pinus patula is
commonly found in mixed forest, associated with other pines and/or angiosperms such as oaks
(Quercus) and also occurs in the Liquidambar forest on mountain slopes in Veracruz that face the
Gulf of Mexico. A wealth of epiphytes, most abundantly on oaks, characterizes these wet forests.

Map MCW-16. Pinus leiophylla var.

leiophylla (red) [n=138, m=126, h=37,
18??–1994] and Pinus leiophylla var. chi-
huahuana (black) [n=113, m=109, h=30,
1846–1995]

The two varieties of Pinus leiophylla

have different distributions, which
only partly overlap. The most wide-
spread variety, var. leiophylla, occurs
from the northern end of the Sierra
Madre Occidental to Oaxaca and
is endemic to Mexico. The more
northern variety, var. chihuahuana,
is widespread in the Sierra Madre
 mexico, central america and the west indies 101

Occidental of Mexico and extends into the southernmost parts of Arizona and New Mexico in
the USA. In the Sierra Madre Occidental, where both taxa are sometimes sympatric, var. leiophylla
is less common. These two taxa have been treated as distinct species, for which there is now some
support coming from DNA based research (Parks et al., 2012). Where the two taxa occur in the
same area, var. chihuahuana tends to occupy lower altitudes with drier forest characterized by
P. cembroides, Juniperus spp. and xerophytes like Opuntia and Arctostaphylos. Morphologically the
two taxa are very similar, especially sharing prolonged cone development and epicormic shoots
not present in other Latin American pines, and show considerable overlap in character states
(Farjon & Styles, 1997). The closer relationship of var. chihuahuana to the next species, P. greggii,
than to var. leiophylla as inferred from the analysis cited above, needs to be tested by genetic research
at the population level, i.e. including several samples from different localities for each taxon.

The two disjunct populations of Pinus greggii have

Map MCW-17. Pinus greggii var. greg-
gii (black) [n=36, m=30, h=29, 1848–1987]
and Pinus greggii var. australis (red) [n=25,
m=25, h=15, 1900–1994]
recently been recognized as distinct varieties. In the
northern part of the species range (Coahuila, Nuevo
León, San Luis Potosí) we find var. greggii and in the
southern part (Querétaro, Hidalgo, N Puebla) var.
australis. Between the two main and more or less
contiguous ranges of the two varieties are two more
isolated localities in San Luis Potosí, Cerro Cam-
panario and a locality in the mountains ca. 5 km N
of Soledad de Zaragoza. The pines at these locali-
ties have been identified as belonging to var. greggii.
This northernmost variety occurs at higher altitudes
(2300–2700 m) and on alkaline soils derived from
limestone and var. australis occurs mostly on acid
(pH 4–5) soils between 1300–2300 m and partly in
mountains with higher rainfall averages. The longer,
more slender and drooping leaves of var. australis
seem to be correlated with higher precipitation, and,
especially on the eastern escarpment of the mountain
ranges along the Hidalgo-Veracruz borderline, with
clouds rising from the Gulf of Mexico. Var. australis is
listed as Endangered (EN) on the IUCN Red List due
to logging, forest clearance and habitat degradation.

Map MCW-18. Pinus lawsonii (black)

[n=67, m=63, h=22, 1894–1994], Pinus
lumholtzii (orange) [n=71, m=66,
h=30, 1850–1991], Pinus pringlei (red)
[n=89, m=87, h=40, 1894–1993]

These three species appear to be

closely related in a recent phy-
logenetic analysis of the genus
Pinus with comprehensive taxon
sampling and using sequence data
from the complete plastid genome
(Parks et al., 2012). The relation-
ship is (P. lawsonii (P. pringlei +
P. lumholtzii)), a clade that is sister
102 mexico, central america and the west indies

to one containing P. patula, P. leiophylla and P. greggii (maps MCW-15 to MCW-17 above). The
ranges of P. lawsonii and P. pringlei are mostly sympatric, but P. lumholtzii occurs to the NW of
these, with no overlapping distribution according to herbarium specimen data. The two species
with sympatric distribution are, according to this result, slightly more distantly related (although
statistical support for this topology was weak) which could perhaps explain their proximity if
they occupy the same habitat. There is certainly overlap here, too, but while the two species
often grow gregariously on relatively dry sites (600–1500 mm precipitation annually), P. pringlei
tends to be more abundant in localities with a higher rainfall (1000–2000 mm), where P. lawsonii
is replaced by other pines. The altitude ranges are also very similar, with P. pringlei distributed
only a few hundred meters higher (1500–2800 m) than P. lawsonii (1300–2600 m). The third spe-
cies, P. lumholtzii, has a similar altitude range as P. pringlei, but it generally receives 500–600 mm
precipitation per year only, much less than the other two. The split between Pinus pringlei and
P. lumholtzii may therefore have involved an adaptation to drier conditions by the latter species,
enabling it to move into the westernmost parts of the Transverse Volcanic Belt and the Sierra
Madre Occidental.

Map MCW-19. Pinus devoniana [n=133, m=125, h=31, 1800–1994]

Pinus devoniana in Oaxaca, Mexico.

Photo credit Aljos Farjon.

Pinus devoniana occurs from about 23° N in Sinaloa, Mexico to the Guatemalan Highlands. It is
more common W of the Continental Divide but occurs on both sides, and becomes more widely
scattered at the northern limits of its range. It has a wide altitudinal range from 700 m to 3000 m
and occurs in pine and pine-oak forests associated with several other species. Its adaptation to
fire through a ‘grass stage’ as a seedling is unique in Mexican pines. Together with P. oocarpa this
species invades recently burnt areas but can then survive a second fire should it occur before it
has developed into a sapling tree. Its large seed cones are not serotinous and open when mature
in the warmth of the sun, so the cones and seeds are unlikely to remain undamaged in a forest
fire. The large seed wings help to transport the seeds from unburnt forest on to the burn. A dry
season from November to May in most of its range favours the fires, but a relatively high annual
precipitation of 1000–1500 mm enables quick restoration of the pines.
 mexico, central america and the west indies 103

Pinus montezumae in Tamaulipas,

Mexico. Photo credit Aljos Farjon.

Map MCW-20. Pinus montezumae [n=159, m=128, h=33, 18??–1994]

Pinus montezumae has a wide distribution, occurring from Nuevo León in NE Mexico and Durango
in W Mexico to the Guatemalan Highlands. This species is most common along the Transverse Vol-
canic Belt and in the Guatemalan Highlands, in most other locations it occurs less frequently among
the pines and oaks that make up much of Mexico’s forests. It is most abundant and best developed
in the temperate zone in central Mexico at ca. 2400–2800 m a.s.l., with annual precipitation exceed-
ing 800 mm. The altitude ranges from 1200 m to 3500 m, with the lowest occurrences in Nayarit and
Nuevo León where this pine touches on the ‘pinyon-juniper’ belt of semi-arid coniferous forest. At
the highest locations of P. montezumae in Nuevo León it is a constituent of cold temperate conifer
forest with Abies vejarii and Pseudotsuga menziesii var. glauca, indicating the wide range of climate
conditions and forest types in which this species can grow even within a short distance.

Map MCW-21. Pinus arizonica var. arizonica (black) [n=57, m=47, h=20, 1874–2004], P. arizonica var. cooperi
(yellow) [n=39, m=37, h=14, 1900–1991], P. arizonica var. stormiae (red) [n=17, m=16, h=11, 1880–1983]

Pinus arizonica has long been considered a variety or subspecies of P. ponderosa, a widely dis-
tributed western North American pine, particularly by US foresters, who knew this taxon from
southern Arizona and New Mexico. However, the greater part of its range is in Mexico where the
distinctions become more apparent. Pinus ponderosa var. scopulorum only just occurs across the
international border in Sonora and Coahuila (map NA-21 on p. 47). Pinus arizonica is a species
of the northern parts of the Sierra Madre Occidental (var. arizonica, var. cooperi) and the Sierra
Madre Oriental (var. arizonica, var. stormiae). Only var. arizonica extends across the border into
Arizona and New Mexico (just) and could occur in Trans-Pecos Texas (no specimens found for
104 mexico, central america and the west indies

our database). There is an isolated population of var. cooperi in the Serranias del Burro in N Coa-
huila; at only 1300 m a.s.l. this is much lower than the common altitude for this variety, well above
2000 m. Not far to the west, in the Sierra del Carmen, is the southernmost stand of P. ponderosa.
A search in this whole region on either side of the international border might yield new localities
for both species. Pinus arizonica occurs in the higher parts of N Mexico surrounded by deserts
(only var. cooperi links up with the pine forests in the southern Sierra Occidental), where rain-
fall is moderate (700–900 mm) and mostly occurs in the winter months. Other pines commonly
associated with it are P. engelmannii and P. strobiformis, but its most common habitat is a mixed
forest of oaks and pines, in which Juniperus can also make an appearance. Pinus arizonica var.
cooperi and var. stormiae are both listed as Vulnerable (VU) on the IUCN Red List.

Map MCW-22. Pinus engelmannii [n=99, m=96, h=23, 1846–1991]

Pinus engelmannii is another N Mexican pine for which the US/Mexican border appears to be an
artificial line; it occurs also in the Chiricahua Mts. and Santa Rita Mts. of SE and S Arizona; the
latter are on the border. These are northernmost outlier populations of a species that is wide-
spread in NW Mexico, mainly in the Sierra Madre Occidental, and occupies scattered localities
in the Sierra Madre Oriental as well. The southernmost localities of Pinus engelmannii are in the
Sierra de Morones of Zacatecas, isolated from the main distribution just like the localities to the
E in the Sierra Madre Oriental. This species occupies a substantial altitude range from 1200 m to
3000 m but is most common between 2000–2500 m, where the pine and pine-oak forests are best
developed. This species appears to be tolerant of snow and frost at these altitudes and the outly-
ing occurrences are possibly remainders of a more widespread range in colder periods during the
ice ages, when the pine forests may have spread across the Altiplano Mexicano.

Map MCW-23. Pinus teocote [n=127,

m=114, h=36, 18??–1993]

Pinus teocote is widespread in

Mexico, ranging from Chihuahua
to Chiapas, yet it is endemic to that
country, neither crossing the US/
Mexican border nor extending into
Central America; the only species of
Mexican Pinus with such an exten-
sive range to be so limited. While
its northern limits could be deter-
mined by climate, it is difficult to
ascertain the causes of its absence
 mexico, central america and the west indies 105

in the countries in Central America that have various species of pines. As was observed by Perry
(1991), who had extensive field experience, Pinus teocote has an extremely wide ecological toler-
ance, indicated by its spatial and altitudinal ranges, the latter from 1000 m to 3300 m. In Tamau-
lipas near the Gulf of Mexico at 1000–1300 m it grows in near tropical conditions, in the Valle de
México and in Puebla at 3000–3300 m it experiences a dry season and temperatures dropping
below freezing in winter. Pinus teocote is most often found in pine-oak woodland of a relatively
open character (no closed canopy) indicating its pioneer position in the succession, depending
on regular disturbances. Its presence on rocky ridges and outcrops or on slopes with thin soil
indicates ‘evasion’ of competition from angiosperms. A conifer species that is capable of a wide
tolerance and survival in poor habitats is likely to become widespread, especially in a dynamic
and degraded environment.

Map MCW-24. Pinus herrerae [n=56, m=54, h=30, 1900–

1994]

Pinus herrerae is another Mexican species

of Pinus mostly occurring W of the Conti-
nental Divide, which means, especially in
the southern half of its range, being sub-
jected to a long dry season from November
to May. From N to S annual precipitation
increases from about 900 mm to 1600 mm
and the seasonality of this rainfall increases
also. Pinus herrerae is a common constitu-
ent of pine-oak forest in which it can occur
sympatric with several other pine species,
at altitudes ranging from 1100 m in the
Sierra de Cuale in Jalisco to 2600 m on
Cerro La Cumbre in the same state.

Map NCW-25. Pinus durangensis [n=55, m=54, h=31,

1897–1991]

Pinus durangensis occurs in most parts of the

Sierra Madre Occidental and in a few locations in
the western parts of the Transverse Volcanic Belt
and adjacent mountains of W Jalisco. This pine is
an important constituent of the ‘yellow pine belt’
in the Sierra Madre Occidental, where it forms
sometimes pure stands but often mixed pine
forest or occurs in pine-oak forest. Similar forests
occur N of the international border in the USA,
but with fewer species of pine, even in Califor-
nia. The variety of pines in this mountain range,
with up to 16 taxa in some areas, is matched or
surpassed (to 18) only in the Transverse Volca-
nic Belt and in the Sierra Madre del Sur (Farjon, 1996). While taxonomic relationships exist with
some pines in the Rocky Mountains, a large number of Mexican pines are not closely related to
these more northern species and probably originated here. However, Pinus durangensis is related
to P. ponderosa and P. arizonica and thus links this area of Mexico with the Rocky Mountains to the
north, as has already been demonstrated for other species in subsection Ponderosae that only occur
in Mexico. The Sierra Madre Occidental is probably a region of fairly recent speciation of the genus
(Farjon, 1996) correlated with Miocene/Pliocene orogeny and Pleistocene climate change.
106 mexico, central america and the west indies

Subgenus Strobus

Map MCW-26. Pinus strobiformis

(black) [n=90, m=83, h=23, 18??–
1992], P. ayacahuite var. ayacahuite
(red) [n=122, m=108, h=40, 1837–
1993], P. ayacahuite var. veitchii (blue)
[n=22, m=20, h=17, 1896–1994]

Pinus strobiformis belongs to a

small group of closely related
species which includes P. flexilis,
P. strobiformis, P. ayacahuite and,
slightly more distant, P. lam-
bertiana. The first three species
form a ‘grade’ in morphological
character states roughly distrib-
uted from N (USA) to S (Hon-
duras). Subspecies and varieties
have been described under the
first three species and some were even raised to species, especially if they occurred more or less
in isolation. Only Pinus strobiformis and Pinus ayacahuite are predominantly Mexican and are
mapped here, the other two species are treated in the chapter North America. Pinus strobiformis
occurs from Arizona and New Mexico to Jalisco and San Luis Potosí, but its limits in both N
and S are not clear cut due to gradation with P. flexilis and P. ayacahuite respectively (Farjon &
Styles, 1997). Pinus ayacahuite var. veitchii has also been classified as P. strobiformis ssp. veitchii
depending on what characters are emphasised. Its distribution falls more or less between the two
species, but more specifically within the range of P. ayacahuite. In the USA, much of what was
formerly identified as P. strobiformis has later been interpreted as P. flexilis var. reflexa, admittedly
another intermediate form between two species. The map presented by Critchfield & Little (1966)
includes both under P. strobiformis, our map includes only those herbarium collections that com-
pare well with that species as known in Mexico. Both species are montane to high montane pines
that can form pure stands but more commonly occur in mixed pine forest or pine-fir forest, less
often and usually at their lowest altitudes (range is ca. 1700–3400 m) in pine-oak forest. Pinus
ayacahuite can grow very tall and become emergent.

Map MCW-27. Pinus cembroides ssp.

cembroides var. cembroides (black)
[n=180, m=157, h=40, 1827–2009], P.
cembroides ssp. cembroides var. bicolor
(yellow) [n=65, m=53, h=26, 1881–1995],
P. cembroides ssp. orizabensis (blue)
[n=25, m=25, h=19, 1900–1986], P. cem-
broides ssp. lagunae (red) [n=18, m=17,
h=14, 1890–1987]
 mexico, central america and the west indies 107

The complex of Pinus cembroides includes

these four taxa, all of which have been
treated as separate species, plus several
other named taxa, most of which are here
treated as synonyms (Farjon, 2001). They
form a clade sister to a clade with P. edu-
lis and P. remota, and this larger clade is
again sister to two other pinyon pines: P.
monophylla and P. quadrifolia (Parks et al.,
2012). There is considerable cross-over into
the USA, but the taxa in this map (MCW-
27) occur mostly or entirely in Mexico. The
Pinus cembroides ssp. cembroides in Hidalgo, Mexico. two varieties, cembroides and bicolor, have
Photo credit Aljos Farjon.
wide distributions that overlap consider-
ably, but with var. bicolor more common in
NW Mexico and S Arizona. The subspecies lagunae and orizabensis have in contrast very limited
ranges; the first is restricted to the higher parts of the Sierra de la Laguna in Baja California, the
second subspecies occurs in a small area in Puebla, Veracruz and Tlaxcala, mostly around Pico
de Orizaba. In northern Mexico and the Southwest of the USA these small pines occur over vast
areas together with several species of Juniperus in more or less open pinyon-juniper woodland,
a formation that occupies the transition zone between desert and montane forest, the latter also
dominated by pines, but much taller ones, or forming a mixed pine-oak forest mostly in Mexico.
The seeds of pinyon pines are relatively large and wingless when detached from the small cones,
and nutritious. The two isolated subspecies occur in oak-pine woodland or, in the case of ssp.
orizabensis, also with Juniperus. Pinus cembroides ssp. lagunae is listed as Vulnerable (VU) and
ssp. orizabensis as Endangered (EN) on the IUCN Red List.

Map MCW-28. Pinus culminicola [n=53, m=50, h=26, 1900–1991]

Pinus culminicola is a shrubby pine which occurs on the

highest ridges and summits of mountains between 3000 m
and 3700 m in a small area in the states of Coahuila and
Nuevo León. This species is closely related to P. cembroides
and can be considered as belonging to that complex of taxa,
but specialized to an existence at high altitudes. Like other
high altitude or high latitude shrubby pines, it forms exten-
sive and often dense thickets excluding other shrubs and
trees. However, at less exposed sites it can be found among
the highest individual trees of species like Pinus hartwegii, Abies vejarii and Pseudotsuga menziesii
var. glauca and in the relative shelter of these it may attain a height of 4–5 m. Pinus culminicola
is mostly growing on rocky, calcareous soils so its limited distribution is probably correlated with
the extent of limestone summit areas in the Sierra Madre Oriental. Fires have proven to be a
menace to this species, which is listed as Endangered (EN) on the IUCN Red List.

Map MCW-29. Pinus remota [n=31, m=29, h=18, 1916–1985]

Pinus remota is another pinyon pine of N Mexico and

SW Texas, but it is uncommon and occurs in isolated
populations on mountains and in rocky canyons at 1200–
1850 m. On the Edwards Plateau in Texas it occurs at
much lower altitude (the type collection is from 450 m).
108 mexico, central america and the west indies

This species appears also to be limited to limestone rock, like P. culminicola, but at considerably
lower elevation and it is a shrub or small tree to 9 m tall that does not form thickets. In Texas
it is commonly associated with junipers ( Juniperus monosperma and J. ashei) and shrubby oaks,
in Mexico it occupies the driest sites where pinyon-juniper woodland is not well developed and
desert plants such as cacti and agaves start to make an appearance.

Pinus nelsonii in the

Sierra El Conche,
Tamaulipas, with AF on
a collecting trip in April
1994. Photo credit Dario
Mehía. This rare pine
is restricted to karst
limestone outcrops in a
mostly volcanic moun-
tain range, the Sierra
Madre Oriental. Many
rare conifers in the
world appear to have
retreated to these soil-
and nutrient-deficient
limestones, where they
do not suffer competi-
tion from faster grow-
ing, large-leaved trees.

Map MCW-30. Pinus nelsonii [n=56, m=45,

h=28, 1898–1994]

Pinus nelsonii is one of the most interesting pines from a taxonomic and phylogenetic point of
view. While sharing diagnostic characters mostly with species in subgenus Strobus, it ‘borrows’
one from subgenus Pinus, i.e. persistent sheaths at the base of needle fascicles. Sometimes con-
sidered to be a pinyon pine, it is not closely related to any of these even in the broadest sense,
but appears to be in a more distant clade both from morphological evidence (Farjon, 1996) and
from recent molecular analyses (Gernandt et al., 2005; Parks et al., 2012). Its closest relatives are
the ‘foxtail pines’ P. balfouriana, P. aristata and P. longaeva, all in the western USA. Pinus nelsonii
has a limited distribution in Coahuila (“Montañas del Carmen”?), Nuevo León, San Luis Potosí
and Tamaulipas, in foothills or lower slopes of the Sierra Madre Oriental. The location in Coa-
huila cited in Perry (1991) is not represented by specimens in institutional herbaria and needs
such confirmation. It is not mapped here for this reason and because there are three mountain
ranges with this name (“Sierra del Carmen”) in that state and we do not know which of these is
meant. All three seem to be too far removed from the main distribution of this species to be a
plausible location. The most extensive populations are found around the Sierra Peña Nevada in
Nuevo León-Tamaulipas. It is restricted to sites on rocky limestone with shallow soils. Its altitu-
dinal range is 1600–2450 m a.s.l. and it is adapted to low and erratic rainfall, mostly in summer
rainstorms, ranging from 300 mm to 600 mm annually. At higher altitudes P. nelsonii may be a
constituent of pinyon-juniper woodland, lower down it is associated with deciduous shrubs and
succulent desert plants such as cacti and yuccas. This species is listed as Endangered (EN) on the
IUCN Red List.
 mexico, central america and the west indies 109

Map MCW-31. Pinus rzedowskii [n=18, m=18, h=19,

1900–1995]

Pinus rzedowskii is another enigmatic Mexican

pine, more closely related to the pinyon pines than
P. nelsonii, but morphologically rather different.
It is extremely rare, known from only a handful
of small populations or sub-populations in close
proximity in the municipality of Coalcomán, W
Michoacán. Recent surveys have discovered sev-
eral additional localities beyond the three that
have been known since this species was discov-
ered in the 1960’s. Not all can be mapped here and
some are so close to earlier known locations that
our geo-referencing based on degrees and min-
utes would not resolve them even if there were
herbarium specimens. All these localities are in
the vicinity of Dos Aguas, a small town or village
whose main economy is based on the exploitation
of the surrounding pine forest for timber. How-
ever, Pinus rzedowskii is not a target species as it
is a small tree not of interest commercially. Most
of its localities are outcrops of karst limestone, as
with P. nelsonii (see previous map and text), often
inaccessible except on foot and that with difficulty.
The real hazard to this species is fire, to which this
pine is not well adapted. Pinus rzedowskii is listed
as Vulnerable (VU) on the IUCN Red List.

Pinus rzedowskii on Cerro de Chiqueritas, an outcrop

of karst limestone in mountains above Dos Aguas,
W Michoacán. The vegetation on these limestone rocks
differs markedly from the surrounding pine forest and
contains other rare species of plants beside this pine.
Photo credit Aljos Farjon.

Map MCW-32. Pinus pinceana [n=70, m=53, h=27,

1837–1994]

Pinus pinceana occurs in Coahuila, Hidalgo,

Querétaro, San Luis Potosí and Zacatecas, mainly
in the Sierra Madre Oriental and its surroundings,
but extending a little further south. Whether this
and the next species, P. maximartinezii, are pinyon
pines depends on how one defines that group or
complex of pine taxa. The two are closely related
and form a clade sister to the pinyon pines in a
narrow (and ‘traditional’) sense. Both differ from
these in some striking morphological charcters,
and from each other. The wingless, edible seeds
are a common feature with ‘true’ pinyon pines, but
we know that this trait, linked to seed dispersal by
110 mexico, central america and the west indies

birds, has evolved independently several times in the genus Pinus, even across the two subgen-
era. The habitat of P. pinceana is in many ways similar to that of P. nelsonii, yet the two species,
although they have partly overlapping ranges at similar altitudes, rarely grow together.

Pinus maximartinezii in the Sierra de Morones W of

Map MCW-33. Pinus maximartinezii [n=16, m=13,
h=20, 1900–2010]
the town of Juchipila, Zacatecas. The pine-oak wood-
land is degraded in many places due to cattle grazing
and too many fires, resulting in a more open land-
scape dominated by older pine trees and scattered
shrubs. Most oak trees have been felled for timber and
firewood. Photo credit Aljos Farjon.

Pinus maximartinezii is one of the rarest pines of Mexico. It is apparently closely related to
P. pinceana (Gernandt et al., 2005; Parks et al., 2012) but morphologically distinct with extremely
large and heavy cones, which are therefore likely to be a recently evolved adaptation. It also
occurs in a different part of Mexico. While known from the Sierra de Morones in S Zacatecas
since its discovery in 1964, recently a second population was found at La Muralla, S Durango, ca.
200 km NW of the first locality. This newly found population is much smaller than the ‘classical’
one, perhaps the main reason why it was not noted earlier; the species with its 1.5–2.5 kg seed
cones is conspicuous enough. In the Sierra de Morones P. maximartinezii occurs at 1750–2400 m
along several km of slopes on the E-side of the mountain, where it is virtually the only pine spe-
cies in mixed pine-oak woodland. Grazing of cattle, too frequent fires and harvesting of seeds for
the market all put pressure on the succesful regeneration of this remarkable pine, which is listed
as Endangered (EN) on the IUCN Red List.

Abies

The genus Abies (Pinaceae) is widely distributed in Mexico and Central America, occurring from
Baja California Norte to Honduras. The presence in Baja California, on the Sierra San Pedro Martír,
is an isolated southern extension of a species widespread in the USA: Abies concolor; it is mapped
and discussed in the chapter North America. Five species are endemic to Mexico: Abies durangen-
sis, A. hickelii, A. hidalgensis, A. religiosa and A. vejarii; only one, A. guatemalensis, extends from
S Mexico into Central America. In several of these species, varieties or subspecies have been
described, but a proposal to reduce the number of species by sinking several of these taxa as
subspecies into A. religiosa based on morphometric studies was published recently (Strandby et
al., 2009). Undoubtedly the taxonomy of Abies in Mexico and Central America needs more work,
 mexico, central america and the west indies 111

including evidence from molecular data. We retain the species and some of the varieties as recog-
nized in Farjon (2010a) in the maps of this Atlas as the separate distributions of most of these can
be used as additional evidence in this research. The genus Abies in this region is exclusively high
montane, with few exceptions ranging from 2000 m to 3550 m a.s.l. and is therefore concurrent
with the high mountains in the region. In times of colder climate during the glacial maxima of the
Pleistocene isolated populations were probably expanding and in contact, only to be separated
again during warmer interglacial periods. This history if correct provides an explanation for the
species complexes taxonomists appear to find today.

Map MCW-34. The distribution of species in the genus Abies in Mexico and Central America.

Map MCW-35. Abies durangensis

var. durangensis (black) [n=29,
m=25, h=11, 1934–1996], A. duran-
gensis var. coahuilensis (orange)
[n=9, m=8, h=6, 1941–1994] and
A. hidalgensis (red) [n=2, m=2,
h=6, 1994]

Abies durangensis var. duran-

gensis is the fir of the Sierra
Madre Occidental, where it
is limited to the western side
of the continental divide and
mostly found on cool N-facing
slopes at altitudes between
1650 m and 3100 m. Most often
it is a constituent of mixed
conifer forest, with several
pines (Pinus) and Pseudotsuga
112 mexico, central america and the west indies

menziesii as co-dominant trees. Abies durangensis var. coahuilensis has been found in two disjunct
populations in the Sierra Madre Oriental, in Coahuila and Nuevo León: the Sierra de la Madera
and to the SE of this location in mountains near General Cepeda and in the Sierra de Arteaga and
Sierra de La Marta. Here the altitude range is 2100–3550 m, higher than in the western range of
this species; at its lower sites it is confined to shady canyons and only on the highest ridges and
summits can it exist on more open slopes, forming dense forests usually mixed with other coni-
fers. This variety is listed as Vulnerable (VU) on the IUCN Red List. Abies hidalgensis is a some-
what doubtful species only known from its type locality, a canyon near the village of Metepec in
Hidalgo; its high number of resin ducts in the leaves seem to link it with A. hickelii, the only other
Mexican fir with high numbers. This species is listed as Vulnerable (VU) on the IUCN Red List on
account of its very limited known distribution.

Map MCW-36. Abies vejarii [n=21, m=20, h=9, 1939–1991]

The three varieties commonly recognized in this spe-

cies are here ignored and mapped under the species
name. Abies vejarii has a distribution partly overlap-
ping with the southern part of the range of A. duran-
gensis var. coahuilensis, but the two species do not
appear to grow together and A. vejarii extends further
south in Nuevo León and Tamaulipas. Its altitudinal
range and ecology are very similar to A. durangensis
so it is interesting that herbarium collections of the
two species examined for this Atlas have been made
in separate localities. Whether the two species are
truly allopatric cannot be decided from these data
and needs additional field observations.

Map MCW-37. Abies hickelii [n=20,

m=19, h=8, 1900–2002]

Abies hickelii has two varieties, here

mapped together under the spe-
cies as they seem to partly overlap
in ranges. This species is restricted
to S Mexico and has been collected
in three areas: Veracruz-Puebla,
Oaxaca and Chiapas (red circle). In
Veracruz it occurs on and around
the volcano Cofre de Perote; not far
from there it is found in the Sierra
de Maxtaloyan in Puebla. Another
relatively extensive distribution is
present in central Oaxaca, but in
Chiapas only one locality is known,
based on a collection by E. L. Little
in 1960 (No. 17959) from “Rancho Nuevo Unidad Sierra Grande, Chajzalar” = Rancho Nuevo (spec-
imen at K). This species occurs at very high altitudes between 2500 m and 3400 m, sometimes
 mexico, central america and the west indies 113

in pure stands on summit ridges but more commonly mixed with high altitude pines e.g. Pinus
montezumae, P. pseudostrobus and P. ayacahuite and with Cupressus lusitanica. Abies hickelii is
listed as Endangered (EN) on the IUCN Red List.

Map MCW-38. Abies religiosa [n=64, m=62, h=18, 1865–1997] Abies religiosa on the Nevado de
Colima, Jalisco, Mexico. Photo
credit Aljos Farjon.

Abies religiosa is the most widespread species of fir in Mexico, even when not including taxa such
as A. hickelii and A. guatemalensis as a subspecies of it, as proposed by Strandby et al. (2009).
However, it is most common and forms extensive populations on the high volcanic peaks of the
Transverse Volcanic Belt, from Jalisco in the west to Veracruz in the east. Well known forests of
Abies religiosa in this region are on Nevado de Colima in Jalisco, Popocatepetl in México and Pico
de Orizaba and Cofre de Perote in Puebla/Veracruz. Further south, in the Sierra Madre del Sur,
several localities are known, but here it is more often mixed with other conifers, mainly pines
(Pinus). The altitude range of Abies religiosa is extensive, too, with lowest occurrences around
1000 m and highest over 4000 m, but most well developed forests dominated by this species are
found between 2000–3400 m. At the highest altitudes it forms either pure stands or is mixed with
Pinus hartwegii.

Map MCW-39. Abies guate-

malensis var. guatemalensis
(black) [n=22, m=20, h=11,
1850–2000], A. guatemalen-
sis var. jaliscana (red) [n=14,
m=14, h=10, 1947–1994]

Abies guatemalensis extends from Jalisco in the west to Honduras in the east; it is here recognized
to have two distinct varieties, A. guatemalensis var. jaliscana (= A. flinckii Rushforth) in Jalisco
and W Michoacán, and A. guatemalensis in Guerrero, Oaxaca, Chiapas, Guatemala, El Salvador
114 mexico, central america and the west indies

and Honduras. On the mountain Nevado de Colima in Jalisco, the firs that occur at lower alti-
tude, around 2300 m and mixed with pines and oaks, are A. guatemalensis var. jaliscana; higher
on the mountain where the fir forest becomes ‘pure’ the species is A. religiosa. The altitude range
of A. guatemalensis var. jaliscana found on herbarium specimen labels is 1830–2450 m. The more
tropical var. guatemalensis generally occurs much higher, to 4100 m in Guatemala and 3800 m in
Chiapas. The highest mountains in Honduras, which are much lower, have small populations on
their summits (Mt. Celaque, Mt. Santa Barbara) at 2750–2820 m. In the tropics, these summit for-
ests are usually mixed conifer forest and are likely to be refuges for cool-temperate conifers such as
Abies, which may disappear in a warming climate, especially when their altitude limits are already
small, as in Honduras. Abies guatemalensis is listed as Endangered (EN) on the IUCN Red List.

Picea

Three species of Picea (Pinaceae) occur in Mexico, all confined to the northern parts of the coun-
try in the Sierra Madres Occidental and Oriental. These mountain ranges are the southernmost
outposts in the western hemisphere of this genus, which is much more widespread and abundant
in Canada and the USA. In Mexico, spruces are confined to cool and moist sites on N-facing slopes
at high altitudes and in shady canyons and only occur in a few scattered localities. One species,
P. engelmanii, is widespread in North America and has a subspecies that is distributed mostly in
Mexico (ssp. mexicana); it is treated in the chapter North America (map NA-39 on p. 57).

Map MCW-40. Picea martinezii [n=8, m=8, h=7, 1982–1997]

Picea martinezii occurs in the Sierra Madre Oriental and is

known from just three areas in Nuevo León, Aramberri, Mon-
temorelos and Zaragoza. These populations were formerly
identified as the similar species P. chihuahuana but belong
to P. martinezii (Ledig et al., 2004). Two localities in the for-
mer have small populations: Cerro Peña Nevada and Cerro El
Viejo. The type locality is ca. 150 km further N at a location
ca. 6 km SE of La Trinidad. Here it occurs in a sheltered can-
yon on limestone at 2100–2200 m a.s.l. in moist forest with
Abies vejarii, Pinus spp. and several species of angiosperm
trees. The smallest population is said to have fewer than
15 mature trees, the largest several hundred. This species is
listed as Endangered (EN) on the IUCN Red List.

Map MCW-41. Picea chihuahuana [n=11, m=11, h=7, 1942–1994]

Picea chihuahuana occurs in the Sierra Madre Occidental,

especially in several localities around the city of Creel in Chi-
huahua and also in Durango. In all there are some 25 locali-
ties with trees of P. chihuahuana known, most with fewer
than 350 trees. Picea chihuahuana occurs in scattered relict
populations on N-facing high mountain sides, often in can-
yons, at elevations between 2150 m and 3400 m a.s.l. In most
stands other conifers, especially pines (Pinus strobiformis and
P. pseudostrobus) and Pseudotsuga menziesii var. glauca are
mixed in and the spruces are unable to dominate the forest,
as they often do on similar moist and acidic mountain soils
in more northern latitudes. The extreme fragmentation of
 mexico, central america and the west indies 115

the global population and logging operations that have indiscriminately taken this spruce with
the other trees are putting this species at high risk of extinction; few populations are within effec-
tive protected areas. Picea chihuahuana is classified as Endangered (EN) on the IUCN Red List.

Cupressus

The genus Cupressus (Cupressaceae), here retained with its non-cladistic circumscription, has
two species in Mexico, C. arizonica and C. lusitanica. Cupessus arizonica and its several varieties
are mainly distributed in the USA although var. arizonica occurs quite far into Mexico; this spe-
cies is mapped and discussed in the chapter North America.

Cupressus lusitanica is a
widespread, high altitude
cypress of mesic environ-
ment, similar to several Asian
species but unlike its conge-
ners in North America, all of
which are adapted to xeric
conditions. The two varieties
differ mainly in foliage and
tree habit, probably partly
influenced by climatic fac-
tors. The variety benthamii,
often treated as a species, is
most common in the cloud
Map MCW-42. Cupressus lusitanica var. lusitanica (black) [n=61, m=51, forests that cover the moun-
h=16, 1700–2002], C. lusitanica var. benthamii (red) [n=23, m=22, h=9, tains near the Gulf of Mexico
1839–2002] in Hidalgo, Veracruz and
Puebla, only a few locations
are known outside this area. Cupressus lusitanica var. lusitanica predominates in the central part
of the Transverse Volcanic Belt and is also common in the Guatemalan Highlands and on the
summits of high mountains in El Salvador, Honduras and even Nicaragua. The species is rare in S
Mexico as well as in N Mexico, so there is a gap in the distribution between central Mexico and
Guatemala, with only a few localities known from herbarium collections. Cupressus lusitanica is
a tall tree which successfully competes with other high altitude conifers and often occurs mixed
with Pinus and Abies, or even forms pure stands allowing almost no sunlight to the forest floor.
The altitudinal range of this species is from ca. 1000 m to nearly 4000 m a.s.l. This species is widely
planted in tropical and subtropical mountains as a forestry tree and has naturalized in several of
the recipient countries.

Juniperus

The genus Juniperus (Cupressaceae) is second only to Pinus in species numbers in Mexico,
17 species are native to this country. The genus occurs across Mexico and into Guatemala
and is present on two islands in the Pacific Ocean off Baja California. Some species naturally
occur on both sides of the US/Mexico border, but mostly Mexico has species endemic to that
country. Species occurring in Mexico but with the greater part of their range in the USA are
mapped and discussed in the chapter North America. These are Juniperus ashei, J. californica,
J. pinchotii and J. scopulorum. A few Mexican species cross the border with Guatemala, but only one,
116 mexico, central america and the west indies

Juniperus standleyi, is mostly Guatemalan, straddling the border with Mexico only at the Volcán
Tacana. All species belong to subgenus Sabina, the junipers with scale leaves in the mature plants
similar to those of Cupressus.

Map MCW-43. The distribution of species in the genus Juniperus in Mexico and Guatemala.

The distribution map of the genus Juniperus in Mexico and Guatemala does not show the same
correlation with the main Mexican mountain ranges as do those of Pinus (map MCW-6 on p. 95)
and Abies (map MCW-34 on p. 111), instead junipers appear to be more scattered. Although most
species are adapted to xeric conditions, their habitat varies from semi-desert grasslands, pinyon-
juniper woodlands, pine-oak forest and subtropical deciduous forest to alpine treeless grassland
or scrubland. They are absent in the deserts proper, but at higher elevations junipers occur in
transitional grasslands, so we find dots on the map in especially the Chihuahuan Desert. They are
notably absent in the Sonoran Desert, both on the mainland and in Baja California; this desert is
the more extreme of the two. Another conspicuous gap is present in the Isthmus of Tehuantepec,
with a wide zone on either side from where Juniperus appears to be absent. This gap is apparent
for other conifers, too, especially the genus Abies (map MCW-34 on p. 111) and some species in
Pinus. The species of both genera on either side of this gap occur at high altitudes and it must be
the absence of such mountains in the gap that explains it. In the tropics, lowlands, unless edaphic
restrictions apply, are often dominated by angiosperm trees. Among Mexican conifers, a few spe-
cies in the Podocarpaceae, Taxus globosa (map MCW-57 on p. 123) and possibly a few pines are
able to compete, but none of the junipers can do so. Tolerance of shade in seedling to mature
phases is the key limiting factor.
 mexico, central america and the west indies 117

Map MCW-44. Juniperus flaccida var. flaccida (black)

[n=38, m=38, h=10, 1800–2008], J. flaccida var. marti-
nezii (red) [n=22, m=22, h=7, 1974–1999], J. flaccida
var. poblana (yellow) [n=19, m=18, h=4, 1804–2006]

Juniperus flaccida has a large distribution through-

Juniperus flaccida var. flaccida in dry grassland in
Oaxaca, Mexico. Photo credit Aljos Farjon.
out much of Mexico, from Big Bend National
Park in Texas, USA to Oaxaca in S Mexico. The
three varieties here recognized differ in their
distributions as follows: var. martinezii is mainly
found in west-central Mexico, var. poblana is in S
Mexico, while var. flaccida occurs through much
of the range of the species as a whole but is not
present in west-central Mexico. The three variet-
ies appear to occur in separate localities and are
found to be sympatric only in a few places, e.g.
Frijol Colorado in Veracruz, where var. flaccida
and var. martinezii were found in the same area
at 2400–2600 m a.s.l. In the northern parts of the
range, J. flaccida var. flaccida is a constituent of
pinyon-juniper woodland, with Pinus cembroides
as the most common pine. In central Mexico and
at higher altitude (the range is 800–2600 m) it
is found in pine-oak forest or woodland, or in
grassland with shrubs of various kinds in more
xeric locations. Juniperus flaccida forms a small
tree to 15 m tall with a short trunk and a broad,
spreading crown, often with drooping foliage.

Map MCW-45. Juniperus angosturana [n=20, m=20, h=12, 1878–1999]

Juniperus angosturana is closely related to J. pinchotii of Texas

(Adams, 2011, p. 83) but occurs in E Mexico, from the Serra-
nias del Burro in Coahuila to Hidalgo. It is a species that has a
limited distribution in the foothills on the W side of the Sierra
Madre Oriental, where it occurs in grasslands, Acacia scrub,
or in vegetation dominated by succulents like Opuntia, Agave
and Yucca, less frequently in pinyon-juniper woodland. It does
not grow taller than ca. 8 m and is often shrubby with a very
short trunk. Due to increasing pressure to use these marginal
grasslands for pasture Juniperus angosturana is now declining
and listed as Vulnerable (VU) on the IUCN Red List.

Map MCW-46. Juniperus coahuilensis [n=19, m=19, h=16, 1886–2006]

Juniperus coahuilensis is related to J. angosturana, J. mono-

sperma and J. pinchotii (Adams, 2011, p. 83), a clade of serrate-
margin scale leaved junipers with one or sometimes two seeds
per cone that is distributed either side of the US/Mexican bor-
der, with J. coahuilensis known mostly from Mexico but also
from a small area in SW Texas, USA. This species is distributed
118 mexico, central america and the west indies

on the interior flanks of the Sierra Madre Occidental and the Sierra Madre Oriental and occurs
in Bouteloua grassland and high desert (1200–2000 m a.s.l.), often in rocky areas or in canyons. Its
capacity to resprout from the base when burnt probably enables this shrubby species to persist
in a grassland habitat.

Map MCW-47. Juniperus saltillensis [n=29, m=25, h=12,

1886–1999]

Juniperus saltillensis is a high altitude species of N

Mexico occurring in Chihuahua, Coahuila, Nuevo
León and Zacatecas. It is a shrub in Bouteloua
grassland with sometimes dense shrubbery patches
dominated by succulents and Acacia, or patches of
Ephedra, mostly on limestone. Less often it is found
in the understorey of pinyon-juniper woodland. This
species is closely related to Juniperus ashei, another
Texan juniper. It is listed as Endangered (EN) on the
IUCN Red List.

Map MCW-48. Juniperus jaliscana [n=9, m=9, h=6, 1944–1987]

This rare species of juniper is only known from two

locations, one in Durango on the border with Sinaloa
near Pueblo Nuevo, the other in Jalisco around Talpa de
Allende, where it occurs in several localities. The precise
locality in Durango, given as “El Puerto de las Viboras,
Bosques de San Francisco” by the collector C. E. Blanco,
who collected it there for Prof. M. Martínez in 1946, has
not been identified or found back, so the dot on our map is
the approximate location. Juniperus jaliscana may be pres-
ent elsewhere between the two known locations, but on
current knowledge it is listed as Endangered (EN) on the
IUCN Red List.

Map MCW-49. Juniperus durangensis [n=27, m=26, h=7,

1887–2000]

Juniperus durangensis is distributed in Aguascali-

entes, Chihuahua, Durango, Jalisco, Sonora and
Zacatecas and is endemic to Mexico. It generally
follows the Sierra Madre Occidental, with some
outlying localities, and according to Adams (2011)
is most often found on rhyolite (an ultramafic rock
type) in openings or clearings of pine-oak forest or
pinyon-juniper woodland. This seems to indicate
 mexico, central america and the west indies 119

that J. durangensis avoids competition from taller trees (it can occasionally grow to a small tree of
6 m but is usually a lower shrub) by adaptation to poor rocky sites within forests or woodland.

Map MCW-50. Juniperus monticola [n=34,

m=33, h=14, 1800–1994]

Juniperus monticola apparently occurs in

two growth forms, a small tree in pinyon-
juniper woodland or other open pine for-
est, and a decumbent shrub at or above
the tree line on the highest mountains.
The latter form is more common and wide-
spread and occurs from the Cerro La Viga
in Coahuila and the Cerro Potosí in Nuevo
León to the high volcanoes in Veracruz.
Cerro Potosí in Nuevo León, Cofre de Per-
ote and Pico de Orizaba in Veracruz and
Nevado de Colima in Jalisco, as well as sev-
eral volcanoes in México (Valle de México)
all have extensive populations of the high
altitude shrub form. The highest altitude
found with collected specimens is 4270 m,
the lowest limit is around 3400 m. Below
this altitude down to ca. 2000 m the tree form appears. The locality in Guatemala shown on the
map, Barillas in the Sierra de los Cuchumatánes at 3250 m is based on a herbarium collection
from the tree form and it occurred in high montane pine forest with Pinus hartwegii. Similarly, the
northernmost point on the map, in W Coahuila near Sierra Mojada, while the growth habit was
not stated on the herbarium label, is likely to represent a small tree or perhaps an erect shrub as
the mountains near that town do not exceed 2500 m. This species occurs at the highest altitudes
attained by conifers on the North American continent.

Map MCW-51. Juniperus blancoi [n=21,

m=21, h=15, 1940–1998]

Juniperus blancoi has three varieties,

but some of these are still imperfectly
known from few specimens and we
map them here together without dis-
tinction under the species. This species
is scattered along the W flank of the
Sierra Madre Occidental from Sonora to
the Transverse Volcanic Belt in widely
separated locations. This may partly
be due to poor recognition and under-
collection of specimens. The species is
described as a small, shrubby tree but
one variety, known from a single local-
ity, the summit area of Cerro Huehu-
ento in Durango, is a low shrub.
120 mexico, central america and the west indies

Map MCW-52. Juniperus deppeana var. deppeana (black) [n=34, m=34, h=9, 1828–2008], J. deppeana var.
robusta (red) [n=18, m=18, h=5, 1908–1986], J. deppeana var. pachyphlaea (yellow) [n=17, m=16, h=6, 1851–
1976], J. deppeana var. sperryi (orange) [n=3, m=3, h=4, 1940–2008] and J. deppeana var. zacatensis (blue)
[n=5, m=5, h=2, 1945–1972]

This variable species has five varieties and is widespread from Arizona and New Mexico to Puebla
and Veracruz. We have mapped the recognized varieties here with different colour dots (orange
and magenta slightly larger) to see if there are distinct distribution patterns for these taxa. There
is considerable overlap, mainly caused by the nominal variety deppeana, which occurs through-
out the range of the species. The far southern populations, disjunct from the remainder, consist
of this variety only. In the northernmost part of the distribution of J. deppeana, the rare var.
sperryi is confined to the USA; its first discovered locality is in W Texas but more recently it has
turned up in Arizona and more localities will likely be found. Var. pachyphlaea occurs mostly
within the USA and has thus far only been found in a few widely separated localities in N Mexico.
Confined to Mexico are var. robusta, widespread in NW Mexico, and var. zacatecensis, which has
only a small extent of occurrence on either side of the borderline between Zacatecas and Dur-
ango. The altitudinal range for the species is from 750 m to 2750 m a.s.l. Juniperus deppeana is an
arborescent shrub or a tree to 25 m tall; some specimen trees in the USA have very large trunks
and are among the largest junipers known. The habitat varies from grassland to pine-oak forest,
with pinyon-juniper woodland probably the most widespread throughout the range at middle
elevations. Juniperus deppeana var. robusta is listed as Vulnerable (VU), var. sperryi as Critically
Endangered (CR) and var. zacatensis as Endangered (EN) on the IUCN Red List.
 mexico, central america and the west indies 121

Juniperus deppeana var. deppeana (Marsha’s Alligator Juniper) in Ari- Juniperus deppeana var. dep-
zona, USA. Photo credit David Thornburg. peana in Puebla, Mexico. Photo
credit Aljos Farjon.

Map MCW-53. Juniperus comitana [n=14, m=14,

h=7, 1942–1995]

Juniperus comitana occurs in Chiapas,

Mexico and in Guatemala. It is an uncom-
mon species growing as a small tree in
rocky places with sparse vegetation at alti-
tudes between 1300 m and 1800 m. Where
it occurs in more accessible oak-pine
woodland the vegetation is often degraded
by excessive wood cutting and overgrazing
and regeneration will be jeopardized. This
species is listed as Endangered (EN) on the
IUCN Red List.

Map MCW-54. Juniperus gamboana [n=18, m=18,

h=8, 1944–2009]

Juniperus gamboana has its main distribu-

tion in Chiapas, Mexico, in Guatemala it has
only been found in Huehuetenango. It occurs
at altitudes between 1800 m and 2200 m in
the cloud zone of montane oak-pine forests,
as evidenced by the abundance of epiphytes,
such as bromeliads, orchids, ferns, mosses and
lichens. In more open pine forest with Pinus
oocarpa it attains 10–12 m, elsewhere in more
rocky terrain it is a stunted tree with a short
trunk. The limited range of this species and
the deforestation observed in Chiapas where
this juniper occurs are causing the population
122 mexico, central america and the west indies

to decline in that state. The situation in Guatemala may be similar but its extent there remains
poorly known. Juniperus gamboana is listed as Endangered (EN) on the IUCN Red List.

Map MCW-55. Juniperus standleyi [n=23, m=18,

h=6, 1876–1973]

Juniperus standleyi is a very rare species known

from Volcán de Tacaná in Chiapas on the bor-
der with Guatemala and from the highlands
in Huehuetenango and in San Marcos in Gua-
temala. It has been most frequently collected
from the Sierra de los Cuchumatánes in Hue-
huetenango, the other locality is on the Guate-
malan side of Volcán de Tacaná in San Marcos.
It is a high altitude tree or shrub often mixed
with Pinus hartwegii between 3000 m and 4240
m, so it occurs almost as high as J. monticola. In Huehuetenango it is also found on limestone
escarpments and rocky slopes, often in grassy scrubland with cultivation of Agave plants for alco-
hol production. Trees have been cut at such levels that on Volcán de Tacaná fewer than 1/3 of
the historical population remains, now mostly of the shrubby form. Sheep grazing in the Sierra
de los Cuchumatánes inhibits regeneration. Juniperus standleyi is listed as Endangered (EN) on
the IUCN Red List.

Taxodium

Map MCW-56. Taxodium
mucronatum [n=61, m=48,
h=13, 18??–2007]

Taxodium mucronatum
is one of two species in
the genus and is almost
entirely confined to Mex-
ico, with just a few locali-
ties in S Texas along the
Rio Grande that forms
the border with Mexico,
and one locality in Gua-
temala. This species is
most common in central
and southern Mexico, but
is found scattered along
rivers NW into Sonora and NE to the Rio Grande valley in Tamaulipas and Coahuila. The Aztecs
had been planting this ‘sacred’ tree which is associated with rain and fresh water centuries before
Europeans conquered their country in the early sixteenth century. The most famous of these
planted trees is the ‘Arbol del Tule’ near Oaxaca, with a fluted trunk 14.5 m at its widest diameter.
Most of the very large trees of this species in Mexico appear to have been planted. Unlike the
botanically very similar T. distichum of the USA this species does not naturally grow in swamps or
lakes, but on the banks of often intermittent streams, from the lowlands to 2700 m a.s.l. Taxodium
mucronatum is a major constituent of gallery woodland or forest bordering rivers, arroyos and
canyon bottoms with more or less perennial water at least near the surface. The presence of T.
mucronatum indeed indicates water, even if only ground water.
 mexico, central america and the west indies 123

Taxodium mucronatum in a dry river bed

in Oaxaca, Mexico. Photo credit Aljos Far-
jon. This species is often considered syn-
onymous with T. distichum in the USA, or
at most to be a mere variety of it. Botanical
characters of leaves, cones etc. are indeed
very similar, but its geographical and eco-
logical distribution as well as its phenology,
are very different. Taxodium mucronatum
is predominantly an upland tree bordering
rivers and intermittent streams. It is semi-
evergreen and produces no pneumato-
phores on its roots and is not tolerant of
sub-zero temperatures (°C) but can survive
long periods of atmospheric drought. It has
perhaps evolved from T. distichum which
may have invaded Mexico during repeated
ice ages when the climate was cooler and
wetter, being selected for drier conditions
during the warmer interglacials.

Taxus

Map MCW-57. Taxus globosa [n=32,

m=29, h=9, 1837–2002]

The genus Taxus is represented

in Mexico and Central America
by a single species, Taxus glo-
bosa. This species occurs in
Mexico predominantly in the
eastern part of the country,
where the dry season is shorter
and mountain forests are moist
much of the year (cloud forest).
Taxus globosa is found from near
Monterrey in Nuevo León to the
Sierra de Juárez in Oaxaca. It has
not been collected in Chiapas,
Mexico but does occur in Guate-
mala, El Salvador and Honduras.
The altitude range of T. globosa is
between 1500 m and 3000 m for most collections, in Guanajuato and Tamaulipas a few have even
been reported lower. There should be suitable habitat in Chiapas, so its absence there is some-
what mysterious if it is not an artefact of collecting activity. Taxus globosa is not a very common
conifer and appears to occur in limited areas long distances apart. In most of its range it is found
as an under-storey tree in high montane cloud forest, in Nuevo León under Abies-Picea forest. In
El Salvador and Honduras the cloud forest community with conifers such as Abies guatemalensis,
Cupressus lusitanica and Taxus globosa is limited to the highest summits around 2500–3000 m.
This species is listed as Endangered (EN) on the IUCN Red List.
124 mexico, central america and the west indies

Podocarpus

The genus Podocarpus is represented in Mexico with three species, P. guatemalensis, P. matudae
and P. oleifolius. The only species with a substantial distribution in Mexico is Podocarpus matu-
dae, which is mapped here. Podocarpus guatemalensis is mapped below in the section on Central
America (map MCW-60 on p. 126) and P. oleifolius (map SA-18 on p. 439) is treated in the chapter
South America.

Map MCW-58. Podocarpus matu-

dae [n=71, m=68, h=17, 1936–2006]

This species occurs from Hon-

duras to Tamaulipas and Jalisco
in Mexico. Podocarpus matu-
dae is found in mixed pine
forest, pine-oak forest, mon-
tane rain forest, and evergreen
cloud forest, with an altitudinal
range from 600 m to 2600 m.
a.s.l. It grows often in ravines
near streams, especially in
drier pine forests in the west-
ern part of Mexico. Most trees
in the broad-leaved forests are oaks (Quercus spp.) but also abundant are other deciduous trees
such as Liquidambar, Magnolia, Ostrya, Clethra, and, especially in Chiapas, Puebla and Veracruz,
species of northern genera like Fagus, Carpinus and Platanus. The rarity of Podocarpus matudae
in Central America, where only three disjunct localities are known from herbarium collections
(the collection from Guatemala is on the border with Chiapas, Mexico) and its relative abundance
is several locations in Mexico, as well as its wide distribution there as the only common species,
seem to indicate that it originated in Mexico. It is probably closely related to P. guatemalensis, a
species with a very wide distribution in Central and South America including parts of S Mexico
(map MCW-60 on p. 126). Due to deforestation in parts of its range this species is listed as Vulner-
able (VU) on the IUCN Red List.
 mexico, central america and the west indies 125

Central America

Geography

Central America as considered here comprises the following countries: Guatemala, Belize, El Sal-
vador, Honduras, Nicaragua, Costa Rica and Panama. From Guatemala to Nicaragua the land is
250–400 km wide, but in Costa Rica and Panama it narrows and these two countries form an
isthmus between the Atlantic and Pacific Oceans which is at its narrowest point in Panama only
ca. 70 km wide. The region is mountainous for much of its length, but with interruptions of low
lying areas mainly in N Guatemala, its Pacific coast, on the Atlantic side of Honduras and Nicara-
gua and in Costa Rica and Panama. This present land bridge between North America and South
America became established in the late Pliocene, ca. 3.5 Ma, earlier there was a Panamanian arc
of islands that enabled ‘island hopping’ for organisms with good dispersal mechanisms.

Conifers in Central America

Families Genera Species Species + infra Countries with Area of occupancy Endemic taxa
conifers in km²
4/8 8/70 24/615 25/794 7/7 10,750/542,750 2
Numbers after ‘/’ are global or continent totals.

Conifers are most abundant in the Guatemalan Highlands, in Belize, Honduras and NW Nica-
ragua. They are absent or nearly absent in the lowlands of Central America, but also scarce to
absent in El Salvador and the southern half of Nicaragua where the country is mountainous.
Much of the density in the more northern parts of Central America is due to species of pines
(Pinus), almost all of which are Mexican species extending to this region, except Pinus carib-
aea var. hondurensis. Other northern conifers such as Abies, Cupressus, Juniperus and Taxus do
not extent further south than Honduras, so those conifers which occur naturally in Costa Rica
and Panama belong to the family Podocarpaceae. Their distribution is scattered in part due to
topography, as all species in the region are montane, but some are limited to a small area of the
mountains in these two countries.
The diversity of conifers in Central America compares with Mexico at the levels of family and
genus, 4/4 and 8/11 respectively. The number of species is much lower (24/80) but this is not
surprising for a much smaller area. More remarkable is the very low number of endemic taxa,
with only two species endemic to Central America: Podocarpus costaricensis and Prumnopitys
standleyi, both confined to Costa Rica. Pinus caribaea var. hondurensis and Juniperus standleyi
are nearly endemic but both taxa have small populations just across the border in Mexico. In
this section we give maps only for the two endemics and Podocarpus guatemalensis which is
most common in Central America. Pinus caribaea var. hondurensis is treated in the next section
of this chapter (West Indies) under the species (map MCW-71 on p. 133) and Juniperus standleyi
has been mapped in the previous section with other junipers of Mexico (map MCW-55 on p. 122).
Most other species occurring in Central America are more widespread in Mexico and are mapped
in that section (see above). Podocarpus magnifolius (map SA-17 on p. 439) and P. oleifolius (map
SA-18 on p. 439) are treated in the chapter South America.
126 mexico, central america and the west indies

Map MCW-59. The distribution of conifer species in Central America.

Map MCW-60. Podocarpus guatema­

lensis [n=61, m=57, h=16, 1922–2008]

Podocarpus guatemalensis has a

very extensive distribution from
Oaxaca in Mexico to N Venezu-
ela, Colombia and Ecuador. It is
most commonly found in Central
America. This species grows to a
large tree 20–35 m tall and 1–1.5 m
d.b.h. in tropical lowland to mon-
tane forest. In the N of its range it
occurs often along streams in pine
forest, e.g. with Pinus caribaea var.
hondurensis or P. oocarpa, but in
Costa Rica, Panama and South
America it is a constituent of evergreen broad-leaved tropical rainforest and becomes a canopy
tree, apparently successfully competing with other trees and growing to similar size. The altitude
range is considerable, from near sea level (in Guatemala and Panama) to 2300 m (in Guatemala)
and 2615 m (in Ecuador) based on herbarium collection data. Podocarp trees of these dimensions
provide valuable timber of high quality and logging has undoubtedly impacted the occurrence
of this slow growing tree. Its great extent of occurrence and its presence in many forest reserves
have kept it, so far, on the safe side of an IUCN category of threat.
 mexico, central america and the west indies 127

Map MCW-61. Podocarpus costari-

censis [n=9, m=8, h=6, 1975–1997]

This species is limited to four

localities in Tarrazu, Costa Rica
within an extent of occurrence
(EOO) of less than 100 km². It
occurs in rainforest at altitudes
from 70 m to 1650 m (based on
herbarium label data) but little is
known of its ecology. It is likely
to be rare or at least uncommon,
as few collections were made in
a country that is botanically well
surveyed. We do not know but can suspect decline due to deforestation at lower altitudes in less
steep terrain and the fact that none of the known localities is in a protected area. Podocarpus
costaricensis is listed as Critically Endangered (CR) on the IUCN Red List. A forest reserve to
protect this species is urgently needed, following surveys into its EOO and abundance, as well as
ecological aspects.

Map MCW-62. Prumnopitys standleyi

[n=17, m=17, h=7, 1888–2000]

Prumnopitys standleyi is the only

representative of its genus in
Central America (for the genus
see map SA-5 on p. 432) and has a
distribution limited to four prov-
inces in Costa Rica. Many herbar-
ium collections have been made
along a stretch of the Pan Ameri-
can Highway that leads alongside
or through a national park, a situ-
ation that leads to the possibility
that this species is more common elsewhere than the map suggests. On the other hand some
surveys have found fewer than one mature tree per hectare, so it is probably rare within its for-
est habitat. Prumnopitys standleyi is a climax forest species in montane tropical rain forest at
altitudes between 1800 m and 3000 m. Due to logging in unprotected areas a serious decline has
occurred and this species is listed as Endangered (EN) on the IUCN Red List.
128 mexico, central america and the west indies

The West Indies

Geography

The West Indies in this chapter comprise the islands in the Caribbean Sea plus Bermuda, an
island in the Atlantic Ocean ca. 1100 km E of the American coast. The Caribbean islands are com-
monly divided into the Greater Antilles and the Lesser Antilles and separately, the Bahamas. The
Greater Antilles are Cuba, Jamaica, Hispaniola and Puerto Rico, with smaller satellite islands.
The Lesser Antilles are again divided into the Leeward Islands and the Windward Islands and
form an arc of small islands from the Virgin Islands E of Puerto Rico to Grenada, continued in a
series of islands off the Venezuelan coast westward to Aruba (‘Southern Antilles’). Trinidad and
Tobago are South American continental shelf islands situated off the delta of the Orinoco. The
largest island is Cuba (114,525 km²), followed by Hispaniola (76,190 km²), Jamaica (11,425 km²)
and Puerto Rico (8,960 km²). The mountainous topography of these islands, especially Hispan-
iola, Jamaica and Puerto Rico, is contrasted by the flat and low-lying islands of the Bahamas. The
Lesser Antilles are an arc of volcanoes and are consequently mostly mountainous, except the
‘Southern Antilles’ which are flat and low-lying. The West Indies are almost entirely within the
tropics, but the Bahamas extend N of the Tropic of Cancer and Bermuda is situated at 32° 18’ N
64° 47’ W. The Caribbean Plate extends from Central America to the Lesser Antilles which are
on its eastern edge; its northern boundary is determined by the Cayman Trench and the Puerto
Rico Trench. Cuba and the Bahamas lie on the North American Plate. Subduction of the Atlantic
Plate under the Caribbean Plate threw up the island arc of the Lesser Antilles, forming a step-
ping stone connection of biogeographical significance between the Greater Antilles and South
America. The climate in the region is tropical monsoonal, with infrequent but violent hurricanes
moving westwards from the Atlantic Ocean.

Conifers in the West Indies

Families Genera Species Species + infra Countries with Area of occupancy Endemic taxa
conifers in km²
3/8 3/70 16/615 20/794 13/24 5,025/238,112 20
Numbers after ‘/’ are global or continent totals.

The distribution of conifer species in the West Indies shows a distinct pattern. In Cuba, conifers
are concentrated in the eastern and western extremities of the island, including Isla de la Juven-
tud (Isla de Pinos). While conifers are more evenly spread over Jamaica and Hispaniola, on the
smaller island of Puerto Rico we see the same E-W division as in Cuba. In the Bahamas there is
also a gap in conifer distribution between the northernmost islands and the Turks-Caicos Islands.
The isolated occurrence on Bermuda is a single species, Juniperus bermudiana. Conifers are pres-
ent on several of the Lesser Antilles and on Trinidad and Tobago, but absent on the ‘Southern
Antilles’. The diversity of conifers in the West Indies is low, with three families, Cupressaceae,
Pinaceae and Podocarpaceae, each represented by a single genus: Juniperus, Pinus and Podocar-
pus respectively. With four species in Juniperus, four species in Pinus and eight species in Podo-
carpus there are 16 species; three additional varieties in Juniperus and one in Pinus take the total
to 20 taxa. However, all of these are endemic to the region. Pinus caribaea has one variety (not
counted here) which occurs outside the region in Central America; it is mapped and discussed in
this section with the two endemic varieties.
 mexico, central america and the west indies 129

Map MCW-63. The distribution of conifer species in the West Indies and Bermuda (not shown here).

Juniperus

The genus Juniperus is present in Cuba, Jamaica, Hispaniola, the Bahamas, Saint Lucia in the
Lesser Antilles and on Bermuda. On Saint Lucia the genus reaches the most southern location in
the Americas at 13° 45’ N (on the mainland around 15° N in Guatemala). This presence seems to
indicate a possible stepping stone migration route between North America and South America
that has either not been realized, or has disappeared due to extinction. Juniperus is absent in
Puerto Rico and all other islands of the Lesser Antilles at present but became extinct on Barba-
dos in historical times. With the inclusion of Bermuda, there are four species present: Juniperus
bermudiana, J. barbadensis (with two varieties), J. gracilior (with three varieties) and J. saxicola.
With the exception of J. barbadensis var. lucayana, all taxa have very small ranges limited to a
single island, sometimes even down to very few living mature plants. This rarity is partly natural
due to restricted habitats, but has in most cases been aggravated by adverse conditions imposed
by humans.

Map MCW-64. Juniperus bermudiana [n=12, m=5,

h=5, 17??–2007]

Juniperus bermudiana is endemic to Bermuda,

a small group of islands in the Atlantic Ocean
ca. 1100 km E of Cape Hatteras on the American
coast at 32° 18’ N 64° 47’ W. These low islands
emerged from the top of a seamount of igne-
ous rocks that arose from the ocean floor to just
below sea level and which is capped by eroded
coralline limestone and aeolian sands derived
from it. These upper sedimentary layers form an atoll, the only one present in the Atlantic Ocean.
Maximum elevation a.s.l. is 30 m and the atoll is but a small part of a now mostly submerged 116
km² large platform. Consequencently, Bermuda is of oceanic origin with no connection to other
lands in its long history. The ancestor of Juniperus bermudiana must have arrived from elsewhere
130 mexico, central america and the west indies

as seed(s) transported by birds (see for other examples in the Atlantic Ocean J. brevifolia, map
EM-39 on p. 167 and J. cedrus, map EM-40, p. 167). Its nearest relative is probably J. barbadensis
var. lucayana, so it may have been transported from the Bahamas (Adams, 2011, p. 79). Juniperus
bermudiana is one of the species in the genus named and described by Linnaeus in 1753, so it was
known to occur in Bermuda long ago. However, its more recent plight has been severe due to the
accidental introduction of ‘scale’ insects to which J. bermudiana, having evolved in isolation, has
no immunity. This epidemic has killed 99% of the trees, while invasive plant species and develop-
ment for leasure tourism have occupied much of the ground once forming its habitat. This species
is listed as Critically Endangered (CR) on the IUCN Red List.

Map MCW-65. Juniperus barbadensis var. lucayana (black) [n=43,

m=42, h=7, 1888–2004], J. barbadensis var. barbadensis (red) [n=6,
m=4, h=4, 17??–1986]

Juniperus barbadensis var. luca­yana in the Blue Mountains of

Jamaica. Here the juniper trees occur between 1300 m and 1600
m in 3–4 very small subpopulations in the same general area
(Parish of St. Andrew). Apart from exploitation, these small
stands could easily be wiped out by a serious forest fire fol-
lowed by rapid succession of coarse grasses and shrubs that
prevent regeneration of the slower growing junipers. Photo
credit Martin Gardner.

Juniperus barbadensis was described and named by Linnaeus in 1753 from Barbados, where it
has become extinct probably during his lifetime. A tiny population was dicovered recently near
the summit of Petit Piton, a volcanic cone on the island of St. Lucia (red circle) in the Lesser
Antilles, Windward Islands. Other populations of junipers in the Bahamas and Greater Antilles
were described as a distinct species, J. lucayana, but comparative studies have shown these to be
mere variants of the species J. barbadensis. The outlying populations on Barbados and St. Lucia
are perhaps the result of chance long distance dispersal. If they are relicts of ‘island hopping’
events this would imply extinction on islands between Hispaniola or Andros Island in the Baha-
mas and St. Lucia. In the absence of fossils we lack evidence for this second possibility. Juniperus
barbadensis var. lucayana is the most widespread of the Caribbean junipers occurring from near
sea level to 1600 m, most often on limestone or coralline sand. It occurs in pine savannas, upland
pine forests or in sparse shrubland on karst limestone. In Cuba, some populations may still be
fairly extensive, but elsewhere this variety of juniper is often down to a few tens of individuals,
due to over-exploitation (wood carving for tourists) and habitat degradation. Juniperus barbaden-
sis var. barbadensis is Critically Endangered (CR) and var. lucayana is listed as Vulnerable (VU)
on the IUCN Red List.
 mexico, central america and the west indies 131

Map MCW-66. Juniperus gracilior var. gra-

cilior (black) [n=11, m=11, h=7, 1887–1986],
J. gracilior var. ekmanii (red) [n=7, m=7,
h=7, 1925–2001] and J. gracilior var. urbani-
ana (yellow) [n=3, m=3, h=7, 1925–2009]

Juniperus gracilior is endemic to the

island of Hispaniola, which divides
into two countries, the Dominican
Republic in the east and Haiti in the
west. Juniperus gracilior var. gracilior
is endemic to the Dominican Republic
and occurs primarily in the Cordillera Central, with outlying populations to the E in the Valle del
Yaque (1700 m) and to the S in the Sierra de Ocoa (1000 m). These two localities are at the high-
est and lowest altitude for this variety. The other two varieties occur on both sides of the border
between the two countries, in the southern part of Hispaniola. Juniperus gracilior var. ekmanii
has been collected at 900–2100 m a.s.l. and occurs in several locations in the Sierra de Baoruco
(Dominican Republic) and in the Massif de la Selle (Haiti). The Morne de la Selle, a mountain in
the Massif de la Selle, also has a tiny population of J. gracilior var. urbaniana. The two are well
separated, with var. ekmanii near Mare Rouge at 1770 m and var. urbaniana near the summit at
2550 m. The population of var. urbaniana in the Dominican Republic is in the Sierra de Baoruco
at Llamdo Savane Perdi (Perdu), E of Charco de la Paloma at 2100 m. Juniperus gracilior var.
gracilior is a tree occurring in moist forest, but its habitat is now mostly degraded to much more
open secondary shrubland. Juniperus gracilior var. ekmanii once grew to very large trees, but of
these only stumps remain (Adams, 2011, p. 197). Juniperus gracilior var. urbaniana in contrast is a
decumbent shrub growing on limestone; in the Sierra de Baoruco the population is in a flat open-
ing surrounded by forest, where it forms spreading patches in grassland. On the Morne de la Selle
only very small and scattered bushes remain due to fires. The species is listed as Endangered (EN),
with var. ekmanii Critically Endangered (CR) on the IUCN Red List.

Map MCW-67. Juniperus saxicola

[n=8, m=7, h=5, 1915–2003]

Juniperus saxicola has a very lim-

ited distribution in the Sierra
Maestra of E Cuba, in the Cordil-
lera del Turquino, mainly on the
E and N ridges leading to Pico
Turquino, at altitudes between
1200 m and 1850 m. This taxon is
a shrubby tree that has retained
juvenile (i.e. needle-like) leaves.
It occurs in the cloud zone of the mountain in low, open forest, often on very steep, rocky slopes
where agaves and sclerophyll shrubs replace larger trees. This species is now listed as Critically
Endangered (CR) on the IUCN Red List.

Pinus

The genus Pinus (Pinaceae) is present in the Bahamas, in Cuba and in Hispaniola. We also include
here Pinus caribaea var. hondurensis of Central America, treating it together with two other vari-
eties of that species which are indigenous in the West Indies. The other species are Pinus tropi-
calis, P. occidentalis and P. cubensis. Compared to Mexico and Central America, and also to the
eastern USA, the diversity of pines in the West Indies is low. Their relationships are with the
132 mexico, central america and the west indies

eastern USA species, so it is likely that the ancestors of the West Indies species migrated from
there to the islands and that subsequent diversification and spread was limited.

Map MCW-68. Pinus tropicalis [n=54,

m=48, h=14, 1856–1990]

Pinus tropicalis occurs in the west-

ernmost parts of Cuba, in Pinar
del Rio and on Isla de la Juventud
(formerly known as Isla de Pinos).
This species is a lowland tree of
pine savannas on sandy soils, with
undergrowth of frequently burn-
ing grasses, dwarf palms and low
shrubs. It is adapted to these conditions by having a ‘grass stage’ as a seedling, developing first
a root system supported by primary leaves on a very short stem. Once this root system is exten-
sive and supplied with hyphens of a mycorrhizal fungus, a stem will quickly grow up to lift the
foliage and first branches above the flames; only then will secondary leaves (true pine needles)
appear and a branched sapling tree develop. This ability to survive frequent grass fires makes
P. tropicalis more competitive than the partly sympatric P. caribaea var. caribaea where such fires
have become too frequent for the latter under human influence. Atlitudinal limits above 250–300
m prevent P. tropicalis from replacing P. caribaea completely. This species is listed as Vulnerable
(VU) on the IUCN Red List.

Map MCW-69. Pinus occidentalis

[n=64, m=55, h=13, 1785–1988]

Pinus occidentalis is an endemic

pine of Hispaniola, reports from
E Cuba rest on misidentifications
(Farjon & Styles, 1997). This spe-
cies is widespread and occurs in
both Haiti and the Dominican
Republic, but both the greater
part of its extent of occurrence and the most extensive stands are in the Dominican Republic.
Pinus occidentalis occurs in diverse habitats from the lowlands at about 200 m a.s.l. to the high-
est mountain ridges (Pico Duarte and Pico La Pelona) on the island at almost 3200 m. Conse-
quently, it occupies a variety of habitats as well as substrates, but the more extensive pine forests
are found in a zone between 900 m and 2700 m where not depleted by over-exploitation. This
exploitation has greatly reduced the area of occupancy and number of trees and several dots on
the map here presented may be based on herbarium collections (their dates span two centuries
from 1785 to 1988) from trees that have since disappeared. Protection measures have proved to
be inadequate in the Dominican Republic but are non-existent in Haiti. This species is now listed
as Endangered (EN) on the IUCN Red List.

Pinus cubensis occurs in E Cuba, from the Sierra Maestra and the Sierra de Nipe through the
mountains to the far eastern end of the island. It is also present in foothills and in ‘pine barrens’
along the coast, so its altitude ranges from ca. 100 m to 1200 m. The coastal plains are alluvial,
but in the hills and mountains these pines are often confined to serpentine rock or soils derived
from this, locally known as Nipe clay. Here very open stands prevail, in contrast to the pine woods
on the alluvium. Pinus cubensis can also colonize disturbed sites and depending on soil may
persist, or on richer soils be later replaced by other trees and shrubs. No pines occur naturally
 mexico, central america and the west indies 133

Map MCW-70. Pinus cubensis [n=65,

m=53, h=14, 1856–1985]

between E Cuba and W Cuba and

in this eastern part P. cubensis is
the only species. The most sub-
stantial mountainous areas are in
the eastern part of the island, but
pines in Cuba are not confined to
higher altitudes with more rain-
fall. Edaphic factors probably play
a more important role than local
climate in the natural distribution
of the genus in Cuba, where areas
with richer soils were, before human interference, densely covered with tropical evergreen angio-
sperm forests.

Map MCW-71. Pinus caribaea var. bahamensis (red) [n=75, m=66, h=14, 1878–2008], P. caribaea var. car-
ibaea (yellow) [n=32, m=26, h=11, 1901–1980], P. caribaea var. hondurensis (black) [n=108, m=102, h=35,
1876–1994]

The three varieties of Pinus caribaea have clearly disjunct distributions and were in the past often
treated as distinct species. One, var. hondurensis, actually occurs in Central America but is here
mapped with the two West Indies varieties. This species is closely related to P. elliottii var. densa
of S Florida, a taxon once considered to be synonymous with P. caribaea. In the Bahamas, var.
bahamensis occurs on Grand Bahama, Great Abaco, New Providence, North and South Andros
Islands and on Great Inagua Island. Further to the E it reappears in the Turks and Caicos Islands
in three distinct subpopulations at Pine Cay, North Caicos and Middle Caicos. These are now all
nearly exterminated by an invasive scale insect that reputedly arrived with ‘christmas trees’ from
the USA. In W Cuba, var. caribaea occupies a small area on Isla de la Juventud and a larger area in
134 mexico, central america and the west indies

Pinar del Río on the main island. It forms pine forests generally above P. tropicalis, but can occur
with that species down to sea level. The most extensive occurence of the species is that of var.
hondurensis, extending from Quintana Roo in S Mexico through Belize, N and E Guatemala, Hon-
duras (including the Islas de la Bahía) and Nicaragua. This is the variety tropical foresters have
become interested in, as it grows well and into a taller tree than the others. Plantations of Pinus
caribaea in tropical South America, Africa and Asia are of this Central American variety, which
tolerates a great variety of soils and a wide range of annual rainfall, in its native area between 660
mm and 4000 mm. Pinus caribaea var. caribaea is listed as Endangered (EN) and var. bahamensis
as Vulnerable (VU) on the IUCN Red List.

Pinus caribaea var. bahamensis in a typical ‘pine key’ in the Bahamas. Photo credit Martin Hamilton.

Podocarpus

Eight species of Podocarpus (Podocarpaceae) occur in the West Indies. All have restricted dis-
tributions, usually confined to one island of the Greater Antilles or a part thereof; one species
occurs across Cuba and another species from Puerto Rico along the string of islands of the Lesser
Antilles. None of these species was included in the phylogenetic analysis of Podocarpaceae by
Biffin et al. (in Turner & Cernusak, eds. 2011).

Map MCW-72. Podocarpus angustifolius [n=20, m=20, h=11, 1859–2004]

 mexico, central america and the west indies 135

Podocarpus angustifolius is endemic to Cuba. It is mainly distributed in the far west and the far
east of the island; an isolated occurrence in the central part of Cuba is at La Forestal Los Chivera
near Piloto, Province Santiago de Cuba. It is quite common in the Sierra de Nipe, Prov. Granma
and futher E this species extends to near Baracoa. In Pinar del Rio in the western part of Cuba
it is more scattered. Podocarpus angustifolius occurs in low rainforest, often on serpentine, at
altidudes between 450 m and 1100 m.

Map MCW-73. Podocarpus coriaceus [n=37,

m=37, h=9, 1879–2006]

Podocarpus coriaceus has a unique distri-

bution pattern in the West Indies, occur-
ring on an arc of mostly volcanic islands
from Puerto Rico to Trinidad. In Puerto
Rico it occurs only in the western (most
abundant) and eastern parts of the island,
at 500–850 m and is absent in the central
parts, where the highest mountain Cerro
de Punta rises to 1338 m. Podocarpus cori-
aceus also occurs on the islands of Saint
Kitts, Nevis, Montserrat, Guadeloupe,
Dominica, Martinique, St. Lucia, Tobago
and Trinidad. Between Puerto Rico and
Saint Kitts and between St. Lucia and
Tobago the islands do not have native
podocarps. The Virgin Islands (UK and USA) are in part mountainous, with the highest point at
530 m. The islands S of St. Lucia such as St. Vincent and Grenada are higher, with summits at 1234
m and 840 m respectively. Podocarpus coriaceus occurs mostly above 500 m but lower altitudes
have been recorded from Nevis (300 m), Tobago (250–500 m) and Trinidad (123–247 m). It seems
therefore that altitude is not the cause of the absence of this species from these islands. Neither
is there a lack of forest. This podocarp seems to have ‘island-hopped’ to arrive at its present distri-
bution (that is at least the most parsimonious hypothesis) and so it probably became extinct not
only on the islands of the Lesser Antilles where it is now absent, but perhaps also in the central
part of Puerto Rico.

Map MCW-74. Podocarpus
ekmanii (black) [n=9, m=9,
h=5, 1910–2004], P. hispanio-
lensis (red) [n=8, m=8, h=3,
1982–2004]

Podocarpus ekmanii is a
rare species with a very
limited distribution in E
Cuba, where it is known
from three mountain
ranges in Holguín Prov-
ince, Sierra de Nipe, Sierra
del Cristal and Sierra de
Moa, and from an isolated occurrence on Loma Azul in Santiago de Cuba Province. Herbarium
collections made by E. L. Ekman and some others early in the 20th century had been left nearly
unrepeated until in 2004 collectors from the Royal Botanic Garden, Edinburgh went back and
collected the species again. Its habitat is described as ‘carrascales’ or rocky ridges and slopes
136 mexico, central america and the west indies

with sclerophyllous trees and shrubs, the rock is either serpentine or limestone and both are
poor in nutrients. The altitude range is between 450 m and 1000 m (on Loma Azul 500–700 m).
Like the surrounding vegetation, P. ekmanii rarely grows taller than 6–8 m and is often shrubby.
Those populations that were revisited appeared to be doing well and under no apparent threat,
the most eastern one in the Sierra de Moa is now in a national park. To the contrary, Podocarpus
hispaniolensis, endemic to Hispaniola (and to the Dominican Republic) is in trouble. It has a
limited distribution in the Cordillera Central where it is found in tropical broad-leaved evergreen
rainforest at altitudes between 750 m and 1200 m. Only five localities are known and in some the
cutting of trees since 1980 has made large trees of this species very rare. Podocarpus hispaniolensis
is listed as Endangered (EN) on the IUCN Red List.

Podocarpus hispaniolensis in the Cordillera Central of the Dominican Republic, the highest mountain range
on the island of Hispaniola. Photo credit Martin Gardner.

Map MCW-75. Podocarpus buchii

[n=14, m=14, h=8, 1914–1981]

Podocarpus buchii is endemic to

Hispaniola and occurs both in the
Dominican Republic and in Haiti.
It is more widespread in Haiti,
where it occurs on limestone
massifs with a lateritic thin soil.
In the Dominican Republic the
species is known from two loca-
tions in montane forest on metamorphic rock types. The forest is mostly evergreen tropical rain-
forest dominated by angiosperms with small leaves on steep, rocky slopes, ridges and summits
at altitudes between 600 m and 2050 m. This species is listed as Endangered (EN) on the IUCN
Red List.
 mexico, central america and the west indies 137

Map MCW-76. Podocarpus purdieanus (black) [n=9,

m=7, h=3, 1843–1987], Podocarpus urbanii (red)
[n=15, m=13, h=5, 1850–2004]

These two species are both endemic to Jamaica

and have very limited distributions. Podocarpus
purdieanus occurs in two separate locations in
the Blue Mountains, in the parishes of St. Ann,
St. Catherine and Trelawny. It occurs in wet
tropical evergreen forest on limestone. The elevation is 800 m a.s.l. at the Mt. Diablo subpopula-
tion; the highest record gives 2000–2500 m on Holly Mount at the crest of the Blue Mountains.
Logging may have caused mature trees to disappear from some of the known localities, while sap-
lings are now also threatened by the cutting of sticks for yam cultivation on the island. Podocar-
pus purdieanus is listed as Endangered (EN) on the IUCN Red List. Podocarpus urbanii has a very
restricted range occurring in the Blue Mountains of E Jamaica, in the parishes of St. Andrew and
St. Thomas. It is a small tree of cloud forest, occurring on ridges or steep rocky slopes between
1200 m and 2400 m altitude. It has recently been assessed as Critically Endangered (CR) for the
IUCN Red List.

Map MCW-77. Podocarpus trinitensis [n=5, m=3,

h=4, 1848–1929]

Podocarpus trinitensis is endemic to Trinidad.

This species is very similar to P. coriaceus
(map MCW-73 on p. 135) and is perhaps only
a shorter-leaved variety of it. In St. Andrew
County the two species have been collected
in the same area and are probably sympatric.
The more western locality on the map is in St.
George County where so far only P. trinitensis
has been collected.
138 mexico, central america and the west indies

Area cladogram of Podocarpus

Podocarpus hispaniolensis
A
Podocarpus purdieanus
Hispaniola species in the West Indies, South
America and Central America.
Jamaica Phylogeny according to Leslie
et al., 2012.

Podocarpus angustifolius Cuba The phylogeny reconstructed

for species of Podocarpus in
Podocarpus urbanii Jamaica the West Indies (6 out of 8
species included in the analy-
Podocarpus costavicensis Costa Rica
sis, Leslie et al., 2012) shows
an interesting biogeographical
Podocarpus sellowii Brazil
pattern. Four species from the
Greater Antilles form a clade
sister to a second clade which
Podocarpus trinitensis Trinidad contains two species from the
B Lesser Antilles (one subclade)
Podocarpus coriaceus
Puerto Rico and two species from Brazil
to Trinidad
and Costa Rica (another sub-
Central-South
clade) plus Podocarpus gua-
Podocarpus guatemalensis
America temalensis from Central and
South America. The clade
containing the above species
is sister to P. matudae from
Podocarpus matudae
Mexico-Central
America
Mexico and Central America
and basal in the entire clado-
gram is P. oleifolius from Cen-
Podocarpus oleifolius
Central-South tral and South America. How
America
can we interpret this phylog-
eny in biogeographical terms?
The species in the Greater Antilles are not closely related to any Central American or South
American species, unlike the two species in the Lesser Antilles. More basal species in the phy-
logeny occur in Central and/or South America, not in the West Indies. The species in the West
Indies appear to be derived from an ancestor (or ancestors) on the mainland. The topology of
this phylogeny indicates as the most likely route the stepping stone arc of islands of the Lesser
Antilles. A relative dating estimate of the two nodes A and B could test this hypothesis. There
are some obvious biogeographical gaps: if P. sellowii and P. costaricensis are indeed most closely
related to P. trinitensis and P. coriaceus the distances seem too great to be plausible. However,
P. costaricensis (Map MCW-61) is possibly a relict species with a past distribution similar to e.g.
P. guatemalensis, and P. sellowii (Map SA-25) may have shrunk from a much greater extent of occur-
rence (EOO) together with its habitat, the Brazilian Atlantic rainforest (Mata Atlantica), which
was much more extensive in wetter periods coinciding with glacial maxima in the Pleistocene.
Europe and the Mediterranean

Geography

In this chapter we include all countries in Europe except Russia and Turkey because the largest
parts of these two countries are in Asia. Also included are countries bordering the Mediterranean
Sea in North Africa and West Asia, except Turkey, and the Azores, Canary Islands and Madeira
in the Atlantic Ocean. Countries in the Caucasus Region and Russia and Turkey are included in
the chapter Mainland Asia and Japan. For completeness the European part of Turkey is included
in map EM-1. Europe as here circumscribed has a total area of 8.75 million km² and the countries
bordering the Mediterranean total 6.33 million km² but most of that vast area is extreme desert.
The total area for this region is 15.08 million km².
Europe is geographically very complex. As a whole it is a subcontinent of Asia (together known
as Eurasia) but in its parts it is made up of large and smaller peninsulas and islands and, sur-
rounded by seas on all sides but the E, Europe is itself a large peninsula. In the N it extends well
within the Arctic Circle with the North Cape at 71° 11’ N and is made up of the large island Iceland
and Scandinavia, mostly a large peninsula but with Finland bordering on Russia and Denmark a
smaller peninsula and islands. Great Britain and Ireland are large islands in the NW nearer the
mainland; small islands lie out in the Atlantic Ocean between Great Britain and Iceland, on the
Mid Atlantic Ridge and W of the African coast. Portugal and Spain make up the large Iberian
Peninsula surrounded by the Atlantic and the Mediterranean, while mainland Italy is a long pen-
insula into the Mediterranean. Further peninsulas and islands mark this largest of the world’s
inland seas. The African coast of this sea is mostly a narrow band or strip of mountains and hills
beyond which stretches the Sahara. Europe has numerous mountains, the highest of which are
the Alps culminating in Mont Blanc at 4807 m. Extensive mountain ranges are also present in
Norway and Sweden, the Balkans, E Europe (Carpathian Mts.), on the border of France and Spain
(Pyrenees) and in Italy (Apennines). Smaller mountain systems exist in Scotland, Wales, central
France, Spain, the large Mediterranean Islands and Lebanon. In North Africa the Atlas Mountains
extend through much of Morocco and N Algeria. Extensive lowlands are situated in NW Europe
including Great Britain and Ireland, in countries around the Baltic Sea and in E Europe bordering
Russia. Although influenced by the ameliorating seas in most parts, the climate of Europe ranges
from subarctic to Mediterranean and from continental to oceanic. In North Africa, mountains and
the Mediterranean Sea temper the aridity and heat of the Saharan Desert. As a result, extremes of
climate in Europe and the Mediterranean are rare and limited to relatively small areas, e.g. alpine
summits and the far N of Finland, Sweden and Norway for low temperatures and some parts of
the N African coast for aridity. Extensive glaciations spreading from Scandinavia in the Pleisto-
cene caused arctic and subarctic conditions in Europe N of the Pyrenees and Alps. The E-W ori-
entation and massive glaciations in these mountain ranges blocked southward migration of trees
in particular, as was possible in Asia and North America. The roughly one million year period of
glaciations alternating with shorter warm intervals has had a profound negative influence on the
tree flora of Europe. Extinctions devastated this flora, including conifers. Recently, many of these
140 europe and the mediterranean

have been reintroduced by people and are doing well in much of Europe, but this Atlas is only
concerned with the native conifer flora.

Conifers in Europe and the Mediterranean

Families Genera Species Species + infra Countries with Area of occupancy Endemic taxa
conifers in km²
3/8 9/70 41/615 61/794 52/53 51,575/15,080,000 43
Numbers after ‘/’ are global or continent totals.

The diversity of native conifers in Europe and the Mediterranean is therefore low, with three
families, only nine genera and 41 species present. (The SW Pacific island of New Caledonia has
43 species.) There is a relatively high proportion of infra-specific taxa but this is to some extent due
to taxonomic splitting, a result of long-time studies of the species involved. All countries except
Vatican City have native conifers, but in many there are only 1–3 species, e.g. Iceland with one,
Ireland with two and the United Kingdom with three. Endemism is relatively high at 70% of taxa
due to barriers such as the sea and the desert that surround the region on all sides but the E where
taxonomic diversity of conifers is low for much of the length of this border. The main corridors out
of Europe and the Mediterranean for conifer species are through Turkey and the Caucasus.

Map EM-1. Distribution of conifer species in Europe and the Mediterranean.

The distribution of species shows clear patterns with areas of diversity versus areas lacking coni-
fer species. The data do not allow us to show the ubiquity of conifers, but with very few species,
in Scandinavia. As with other general distribution maps that include the northern boreal for-
ests, there are not enough herbarium collections to cover this. Similarly in Great Britain, Ireland
and Iceland, which have three and only one native species of conifer respectively, there is but a
sampling of localities represented. In the rest of Europe and the Mediterranean, the distribution
 europe and the mediterranean 141

of conifer species is more accurately represented by herbarium collections. Conifers are clearly
concentrated in the mountains and rare or even absent in the broad lowlands. The Atlas Moun-
tains, the highlands of Spain, the Pyrenees, the Alps from Nice to Vienna and adjacent mountains
such as the Jura, Vosges, Black Forest and Bavarian Forest, the Carpathians and the mountains
in the Balkans, as well as Corsica, Crete, Cyprus and furthest E the Lebanon all stand out on the
map. In contrast, much of France including the high Massif Central, England and Ireland and the
lowland plain from Belgium to Latvia are nearly devoid of native conifers. Juniperus communis
and, locally as in England, Taxus baccata, account for all the dots on the map in these regions.
Other conifer-poor regions are the southern interior of Spain and adjacent Portugal, parts of Italy,
Hungary and the lower Danube plains in Bulgaria and Romania. The empty area in Spain is too
dry and the plains of Hungary and the lower Danube are (originally) grasslands. Poor availability
of herbarium collections, not absence of conifers, accounts for the few dots in Serbia. To the E,
lowland forests of birch and oak and (former) steppes make up much of the landscape in Belarus
and Ukraine and it is not until deep into Russia that conifers become again abundant.

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Map EM-2. Map of Europe and the Mediterranean showing numbers of conifer genera per 0.5 degree cell.

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Map EM-3. Map of Europe and the Mediterranean showing numbers of conifer species + infraspecific taxa
per 0.5 degree cell.
142 europe and the mediterranean

Map EM-4. Rarity values per 0.5 degree cell. Cell score is the total of the rarity calculations for each species
in the cell. The rarity calculation is 1 divided by the total of cells occupied by a species. Thus, a species that
occurs in one cell scores 1 while a species that occurs in 10 cells scores only 0.1.

The diversity maps EM-2 and EM-3 show a similar pattern with highest diversity concentrated in
the Alps. Conifer diversity increases in Europe fron N to S but it remains low overall, with most
degree cells not exceeding 4 genera and/or 7 species + infraspecific taxa. Outside three cells in the
Alps, no 0.5 degree cell contains more than seven species. At genus rank there is some moderate
diversity also in the Carpathian Mountains, in the Balkans, in Spain and in the Atlas Mountains of
North Africa. Species numbers are moderately high in some Mediterranean parts of Spain, in Cor-
sica, the S Balkans and the Crimea. The pattern of conifer distribution in map EM-1 is confirmed
in these diversity maps, but with exceptions, e.g. the Pyrenees being much less diverse than the
Alps. Pleistocene refugia for most European trees were mainly in the E and SE and in S Spain
(e.g. Walter & Straka, 1970) and the Pyrenees became isolated from several formerly widespread
conifer taxa e.g. Larix decidua, Picea abies, Pinus cembra and P. mugo which returned no further
W than the Alps after the most recent glacial maximum. Other species, if not reduced to extinc-
tion in Europe, failed to expand again and are confined to their last refugia, e.g. Abies pinsapo in
S Spain and N Morocco and Picea omorika in a small area in Bosnia and Serbia. The latter species
was common in Central and NW Europe during earlier interglacials. Map EM-4 gives the distribu-
tion of more or less narrow endemics. The 0.5 degree cells with the highest values all point at local
endemics: Abies pinsapo var. marocana in N Morocco, A. numidica in N Algeria, A. nebrodensis in
Sicily, Picea omorika in Bosnia and Serbia. Taxa with a wider but limited distribution are found
in Cyprus and in the Algerian Sahara. Northern and Central Europe have no endemic conifers. In
part this pattern is congruent with other boreal forest regions, but it also reflects the high rates
of extinction suffered by trees N of the Alps and Pyrenees during the Pleistocene, leaving only a
few successful colonizers to return and spread widely.

Pinaceae

The family Pinaceae is represented by four genera, Pinus, Picea, Larix and Abies and is the most
diverse group with a total of 25 species, more than half of all species in the region. In Scandinavia
Picea abies and Pinus sylvestris together form the taiga forest, in some parts mixed, elsewhere one
of these is the only conifer tree present. In Great Britain and Ireland the only native pine popula-
tion is in the Highlands of Scotland, again P. sylvestris. Abies and Larix appear in central Europe,
 europe and the mediterranean 143

where also a few more species of Pinus are present, but it is only in the Balkans and the Mediter-
ranean that diversity of species in this family rises to make up the total mentioned above.

Pinus

The genus Pinus has 13 species in Europe and the Mediterranean (including a species in the
Canary Islands). Several of these have been subdivided into subspecies and/or varieties, and sev-
eral of these have again been raised to species. This is not the place to discuss taxonomy and the
Atlas follows Farjon (2010a) for subspecies under P. mugo, P. nigra and P. pinaster and varieties
under P. brutia. The Balkans and the Mediterranean have the greatest of this limited diversity,
in part reflecting the refugia for tree species that existed S of the Alps and Pyrenees during the
Pleistocene glaciations.

Map EM-5. Pinus sylvestris [n=327, m-308, h=40, 1587–2009]

Although the largest part of the enormous range of Pinus sylvestris is outside Europe, we map
it here as it is one of the species described by Linnaeus from N Europe (probably Sweden) and
our data for Europe are much more complete than for Asia. Numerous varieties (or subspecies)
have been proposed and two (var. hamata and var. mongolica) in addition to var. sylvestris are
recognized in Farjon (2010a) but not separated on this map. Pinus sylvestris occurs from Portugal
in the W to Khabarovskiy Kray, Russian Far East, i.e. across Eurasia for a distance of 10,000 km.
Its northernmost population is in Finnmark, Norway at nearly 70° N and in the Sierra Nevada
of Spain it reaches furthest S at 37° N. Apart from the Sierra Nevada in Spain and another dis-
junct population in the Scottish Highlands, P. sylvestris in Europe outside Scandinavia follows the
mountains from N Portugal and Spain to E Europe and the Balkans. Occurrences in the lowlands
of W Europe outside Sweden and the Baltic coast are all naturalized from plantations and not
native, or very doubtfully native e.g. in the margins of (former) blanket bogs on the North German
Plain. This species is very common e.g. in the Netherlands in lowland heath and sand dunes, but
only since plantings of pines on these ‘wastelands’ took off in the 19th century. As studies of suc-
cession in lowland forests and woods in W Europe have shown, even on nutrient-poor sandy soils
the ‘naturalized’ pines will eventually give way to oak and beech forest. Natural and persistent
lowland occurrences begin to appear in E Germany, Poland and the Baltic States and continue
further E as the climate becomes increasingly continental. The natural W limit of P. sylvestris in
the Central European lowlands coincides with the 0–+1° C average January temperature isotherm
(Ellenberg, 1988). Pinus sylvestris is also common in Turkey and eastward as far as the Caspian Sea
and on the S-side of the Caucasus (var. hamata in part). In Siberia this species is widespread, but
we lack enough herbarium collections to show this. In Mongolia and NE China var. mongolica is
recognized, but the distinction with var. sylvestris is not well established and specimens from the
Russian Far East are usually classified as var. sylvestris.
144 europe and the mediterranean

Map EM-6. Pinus mugo ssp.

mugo (black) [n=112, m=109,
h=28, 17??–2008]; P. mugo ssp.
rotundata (red) [n=45, m=45,
h=8, 1827–2011]; P. ×rhaetica
(yellow) [n=5, m=4, h=4,
1887–1982]

Pinus mugo is a variable species occurring in the mountains of Central Europe including the Alps,
in the Carpathians of Central and Eastern Europe, in the Balkans and in the Abruzzese Moun-
tains of central Italy. In this variable species two subspecies are recognized, also treated as species
within a “Pinus mugo aggregate” (Schmidt, 2011). Pinus mugo ssp. mugo is a montane to subalpine
shrubby pine adapted to deep snow cover and avalanches and occurs exclusively in the moun-
tains, with concentrations in the Alps and in the Carpathians. It often forms dense thickets of
semi-decumbent stems curved down-slope to give least resistance to avalanches and is common
in snowy cirques and on exposed ridges. Its altitude ranges from 600 m to 2700 m, the lowest in
the Tatra Mountains (W Carpathians), the highest in the Abruzzese (Appenines). Pinus mugo ssp.
rotundata occurs mostly in and around peat bogs and its habit from low shrub to upright tree
appears to depend on variations in drainage of the soil, with low shrubs in wet moor habitat. In
the Alps it has been found up to 1800 m but in other mountains of Central Europe it usually occurs
at lower altitudes. Draining of moorland, peat digging and afforestation with Norway Spruce (Picea
abies) have caused this subspecies to be listed as Endangered (EN) on the IUCN Red List. Pinus
×rhaetica is the putative hybrid between P. mugo and P. sylvestris and does indeed have character
states of both. It occurs sporadically in areas where both parents are present, but seems restricted
in the Alps to the drier inner valleys where P. sylvestris is common. It has been reported from the
Carpathians but herbarium specimens were not available to show this on the map.

Pinus mugo ssp. mugo in the

Tatra Mountains, Slovakia. In
former glacial cirques like this,
above the tree line, P. mugo forms
extensive carpet-like thickets on
glacial moraine and scree and
can be completely covered in
snow for several months. Photo
credit Peter Schmidt.
 europe and the mediterranean 145

Map EM-7. Pinus uncinata [n=66,

m=64, h=19, 1832–2007]

Pinus uncinata is closely related to

P. mugo and is sometimes included
in that ‘aggregate’ species. There is
confusion especially with P. mugo
ssp. rotundata in its erect tree
form. Pinus uncinata occurs from
NE Spain to the Bavarian Alps and
is especially widespread in the
Pyrenees. [Pinus mugo does not
occur in NE Spain or in the Pyr-
enees.] Isolated populations are found on Mont Ventoux in S France, in the Vosges Mountains in
E France and near Mertingen in Bavaria, Germany. Occurrence in the Massif Central in France is
probably not natural as it is known that this tree was planted there in the 19th century. In the Alps
of E Switzerland and W Austria it is partly sympatric with P. mugo but occurs mostly on limestone
substrates on slopes and ridges with less snow cover (not in cirques and avalanche pathways).
The altitude range is 1000–2300 m. It is always an upright tree.

Map EM-8. Pinus nigra ssp. nigra (black) [n=70, m=70, h=11, 1843–2009]; P. nigra ssp. salzmannii (purple)
[n=22, m=22, h=12, 1835–2007]; P. nigra ssp. laricio (red) [n=15, m=15, h=4, 1871–2010]; P. nigra ssp. dalmatica
(yellow) [n=6, m=6, h=3, 1938–2003]; P. nigra ssp. pallasiana (blue) [n=60, m=58, h=10, 1853–2008]; P. nigra
ssp. pallasiana var. fastigiata (red triangle) [n=1, m=1, h=1, 2005]

Pinus nigra is a widespread and variable species occurring from Spain in the W to countries
around the Black Sea in the E. It is mainly a Southern European species with extension into
Turkey and some outlying populations in coastal North Africa (Algeria, Morocco), the Crimea
(Ukraine) and the E Black Sea coast. E Austria, from where the species was originally described
(Wiener Wald) is its northernmost outpost. Because the variation appears to be correlated with
distribution it has been recognized in five subspecies (e.g. in Flora Europaea) which are mapped
here. Pinus nigra ssp. nigra is an upland subspecies widespread in the Balkans S to Peleponnisos
in Greece, in E Austria and N Italy, where it occurs in foothills of the Alps, and in E Romania
(Caras Severin). The altitude ranges from 100 m to 2000 m a.s.l. with most of the occurrences
above 1200 m in Greece. It can form extensive pine forests, often mixed with P. sylvestris. On the
Dalmatian coast of Croatia including some of the islands in the Adriatic Sea, occurs a more or less
dwarfed form recognized as ssp. dalmatica. It is threatened by habitat degradation as it occurs
in poor rocky soils where goats are taken to browse the sparse vegetation, including seedlings.
It is listed as Endangered (EN) on the IUCN Red List. The most western occurrences of P. nigra are
classified as ssp. salzmannii. This subspecies occurs in S France (Hérault, Pyrenees), Spain (scat-
146 europe and the mediterranean

tered from the Pyrenees to the Sierra Nevada), Morocco (Rif Mountains) and N Algeria (Djebel
Djurdura). The last two localities (Rif Mountains not mapped) are the only places where Pinus
nigra occurs along the S coast of the Mediterranean Sea. Pinus nigra ssp. laricio has a disjunct
distribution in Corsica (‘Corsican Pine’), in E Sicily and in the mountains of Calabria. This subspe-
cies occurs at altitudes between 950 and 1900 m in mountains with ample precipitation; Sardinia
and most of Sicily are too dry to sustain this pine. Pinus nigra ssp. pallasiana occurs in Turkey
(Anatolia), Cyprus, Ukraine (Crimea) and on the Russian part of the coast of the Black Sea (Kras-
nodarskiy Kray). Its distribution in Turkey is remarkable as it is absent in the most western part
of Anatolia, where mountains with suitable habitat would seem to occur. To the E it is limited by
increasing aridity. In Cyprus it is restricted to the Troödos Mountains. Its altitude range is from
100 m (Crimean coast) to 2100 m and it forms pine forests above a zone with P. brutia where the
two species occur in the same area. A form with erect branching, var. fastigiata, was found in
Kütahaya in W Anatolia in what became the Vakif Pinewood Nature Reserve. Pinus nigra, espe-
cially the Corsican Pine, is extensively used in forestry throughout Europe and beyond. We have
mapped only natural occurrences and excluded all others, including naturalized ones. However,
herbarium labels are not always clear about this and it may be that a few dots on the map, if
within the natural range, are actually based on (former) plantations. A natural hybrid between
P. nigra ssp. nigra and P. sylvestris occurs in Austria and has been described as the nothospecies
P. ×neilreichiana; we have insufficient herbarium based data to map it.

Map EM-9. Pinus pinaster ssp. pinaster

(black) [n=17, m=17, h=8, 1842–2010];
P. pinaster ssp. renoui (red) [n=8,
m=8, h=5, 1861–2008]; P. pinaster ssp.
escarena (yellow) [n=44, m=43, h=12,
1849–2010]

Pinus pinaster occurs in SW Europe

and the W Mediterranean and
three subspecies are recognized.
This species has also been widely
planted, e.g. in what are now
extensive ‘forests’ in Les Landes on
the coast of SW France, and natural occurrence can sometimes be difficult to establish because
it can easily naturalize from these plantations. Although mostly a coastal or near coastal species,
in Spain what are thought to be natural stands occur in the interior as well as on the Mediter-
ranean coast. Pinus pinaster ssp. pinaster is the Atlantic subspecies; some occurrences in interior
Spain are perhaps not originally natural but mapped here for lack of evidence that they are from
planted sources. In France, the extensive plantings of this subspecies in Les Landes are excluded.
The most widespread subspecies is P. pinaster ssp. escarena which occurs mostly along the Medi-
terranean coast from Malaga in S Spain to Messina in Sicily, but with a few localities in Spain
and S France away from the coast. Pinus pinaster ssp. renoui occurs along the Mediterranean in
N Morocco and two localities far to the E from there inAlgeria and Tunisia. This subspecies is
listed as Endangered (EN) on the IUCN Red List. Along the Mediterranean P. pinaster occurs on
sandy coastal dunes or sometimes higher rocky hills, often with P. halepensis and P. pinea, or in
evergreen oak woodland or even maquis. This species is frost sensitive, which is probably a reason
why only in the far S of its range it ascends into the mountains. In Morocco it reaches 2000 m a.s.l.
and is a constituent of mixed coniferous forest with Pinus nigra ssp. salzmannii, Abies pinsapo var.
marocana, Cedrus atlantica, and Taxus baccata. Common angiosperm trees in this forest type are
Quercus ilex and at the higher altitudes Populus tremula.
 europe and the mediterranean 147

Map EM-10. Pinus pinea [n=50, m=48, h=16, 1599–2007]

Pinus pinea is an Atlantic-Mediterranean species with a mainly coastal occurrence. It is most

abundant in S Portugal and S Spain, where it may also occur naturally at some distance from the
sea. The extent of its natural distribution is uncertain; although two observations may be correct:
no natural populations are known from the S coast of the Mediterranean and the abundance and
natural regeneration on the coastal sand dunes and flats in the S of the Iberian Peninsula appear
to indicate that here at least it is indigenous. The Romans planted this pine for its edible seeds,
which are still marketed today. Old stands of this pine in e.g. Italy, Greece and Turkey although
usually not on coastal dunes have a natural undergrowth of maquis and the pine trees regenerate
there, too. It is possible that P. pinea is also indigenous there and that its natural range extends
as far E as the Black Sea. Any obvious plantings in towns and cities, or occurrences far from the
coast, or outside what seems the climatic limits for natural regeneration, have been excluded on
the map. It will probably be impossible after a history of more than 20 centuries of planting to
definitely establish the natural distribution of this iconic Mediterranean pine.

Pinus pinea on the Algarve coast, W of Faro, Portugal.

Photo credit Aljos Farjon.

Map EM-11. Pinus heldreichii [n=50, m=49, h=14,

1888–2008]

This species occurs in the Balkans, with an out-

lying population in S Italy. More specifically,
P. heldreichii occurs in S Croatia (Mt. Orje),
Bosnia-Herzegovina, Montenegro, S Serbia
(Kosovo), Albania, SE Macedonia, W Bulgaria
(Rila and Pirin Mts.), N Greece and Thessalia
(Mt. Zygos) and in Basilicata (Mt. Pollino) and
Calabria (Mt. La Montea) in S Italy. Pinus hel-
dreichii is according to molecular evidence not
closely related to P. nigra but to a group of Mediterranean pines including P. brutia and P. pinea,
which are in turn related to P. canariensis and the Asian species P. roxburghii (Parks et al., 2012).
148 europe and the mediterranean

Pinus heldreichii belongs to a group of pines which may have spread along the belt of mountains
bordering the remnants of the Tethys Ocean during the Neogene. Its occurrence is scattered, with
small populations often growing at high altitudes (2200–2640 m) on rocky limestone crags or
steep scree slopes. It is slow growing and some trees in Calabria are believed to be well over 1000
years old. In the Balkans this species is planted in forestry and on better soils at lower elevation
it grows much more rapidly than in its severe natural habitat.

Map EM-12. Pinus peuce [n=39, m=39,

h=10, 1839–2008]

Pinus peuce is another Balkan spe-

cies; it occurs in Serbia, Monte­
negro, Albania, Macedonia (and
on the border with or just inside
Greece) and Bulgaria. Extensive for­
ests dominated by this pine are
uncommon but occur in Bulgaria
in several places in the Rila and
Pirin Mountains and in Macedo-
nia in the southern Sar Mountains
from Mt. Pelister to the border with Greece. It is also often associated with Picea abies and Abies
alba or A. borisii-regis, the latter in Bulgaria. This species appears to avoid limestone and occurs
on silicate-rich rocks and serpentine. Its altitude range is 600–2200 m. Pinus peuce is probably a
Tertiary relict that has survived severe contractions of its range due to Alpine glaciations during
the Pleistocene. Fossils dating from before the Pleistocene possibly belonging to this species have
been found N of the Alps.

Map EM-13. Pinus cembra [n=68, m=67, h=16, 1870–2008]

Pinus cembra is widespread in the Alps and occurs in more isolated localities in the Carpathian
Mountains. In the Alps it is found from the Alpes Maritimes in France to the Julian Alps in
N Slovenia (not shown on map: no herbarium data) and Steiermark in Austria. In the Carpathians
it occurs in the High Tatra Mountains on either side of the border between Poland and Slovakia, in
a few small populations in W Ukraine, and in the Munti Bucegi in Romania. W of the Carpathians
in Romania it is known from a few localities, the only one mapped is Mt. Retezat. Pinus cembra is
a subalpine species forming pure stands or mixed with Larix decidua on slopes, glacial moraines
and ridges above 2000 m to a maximum of 2600 m. At lower altitude, down to ca. 1000 m it is only
a minor component in conifer forest dominated by Picea abies. Pinus cembra forms a mutualistic
relationship with the corvid bird Nucifraga caryocatactes which feeds on its large seeds and stores
 europe and the mediterranean 149

them away for later retrieval to feed its young. Many of these seeds germinate after being carried
by the bird upslope, expanding the pine forest into the increasingly abandoned summer pasture,
and by this spreading slowly restoring the artificially lowered tree line.

Map EM-14. Pinus halepensis [n=163, m=157, h=17, 1842–2009]

Pinus halepensis is very widely distributed in the Mediterranean region including Portugal. It is
mostly a coastal species but does occur in inland areas of Morocco, Algeria, Tunisia, Spain, Lybia
and Jordan which have similar climatic conditions, fire frequencies and vegetation. Along the
European shores of the Mediterranean P. halepensis occurs as far E as Greece. On the Aegean
islands, as well as in Crete, Cyprus and Turkey it is replaced by the very similar species P. brutia.
The most eastern area for P. halepensis is the far E coast of the Mediterranean, where it is pres-
ent in Lebanon, Israel, the West Bank, and Jordan. In much of North Africa its occurrence is very
disjunct because the Sahara borders the Mediterranean Sea almost continuously from the Nile
Delta in Egypt to the Gulf of Gabès in Tunisia, only allowing this species to exist in Appolonia
and Tripolitania. Pinus halepensis occurs in fire-prone vegetation (maquis, garrigue) but can form
dense stands with diminished undergrowth when fires have been absent for a longer time. It does
not ascend into the mountains except in S Morocco where it reaches 1700 m a.s.l. It is likely that
increased fire frequency caused by human activity has helped this pine to spread where formerly
oak woodland would have prevailed.

Map EM-15. Pinus brutia var. brutia (black) [n=93, m=85, h=12, 1825–2009]; P. brutia var. eldarica (red)
[n=22, m=20, h=8, 1903–2008]; P. brutia var. pityusa (yellow) [n=21, m=21, h=7, 1892–2002]; P. brutia var.
pendulifolia (red triangle) [n=1, m=1, h=1, 1989]
150 europe and the mediterranean

Pinus brutia occurs mostly to the E of the area considered in this chapter, but we present the map
here because this species is not only closely related to P. halepensis but often confused with it.
There is only some overlap in their distribution in parts of Greece. Apart from a few localities in
Peloponnisos and on Thasos to the W, P. brutia var. brutia occurs in Crete, on Thira, Karpathos,
Rhodos and other Greek islands along the Turkish coast. Both species are absent or very rare in
the central Aegean islands. Pinus brutia var. brutia also occurs in Cyprus and is widespread in
Anatolia, especially in the far W and in the S, less common in the N along the Black Sea coast of
Turkey and absent in most of the interior. Pinus brutia var. eldarica occurs in two areas far to the
E, in the mountains of Kurdistan in Iraq and adjacent Iran (probably also Turkey but no herbarium
data) and in foothills of the Lesser Caucasus in Azerbaijan and Georgia. Pinus brutia var. pityusa is
distributed along the NE shore of the Black Sea from the Crimea to Georgia. It is the only variety to
be threatened and is listed as Vulnerable (VU) on the IUCN Red List. Pinus brutia var. pendulifolia
is known from one locality on the S coast of Turkey. In proximity of the Mediterranean Sea and the
Black Sea P. brutia can form extensive pine forests, sometimes mixed with Cupressus sempervirens
and Juniperus excelsa, and invades maquis vegetation when fire has been absent for several years.
Similar to P. halepensis its distribution coincides with the Mediterranean climate zone and it is
replaced by other conifers or angiosperms at altitudes above 1000–1500 m where summer precipi-
tation increases. Pinus brutia var. eldarica in Iraq occurs in a more arid region at 800–1200 m a.s.l.
and the southernmost populations of P. brutia in Cyprus occur between 1100 m and 1550 m.

Map EM-16. Pinus canariensis [n=24, m=24,

h=7, 1827–2009]

Pinus canariensis is an endemic of the

Canary Islands. It occurs on Tenerife,
Gran Canaria, La Palma and Hierro.
The two islands nearer the African
coast, Lanzarote and Fuerteventura,
are too dry for this pine, but it is also
absent on Gomera, which is not. Pinus canariensis forms extensive forests on Tenerife on slopes
surrounding the old caldera in the centre of the island, where it has colonized old lava flows and
fields of scoria. Few other trees are present on this forbidding substrate but eventually a shrub layer
develops under the often sparsely distributed pine trees. Pinus canariensis is an interesting species
biogeographically. Its closest relatives in a phylogenetic context are P. roxburghii (W Himalayas) and
P. pinea (Mediterranean), followed by P. pinaster (Iberian peninsula, W Mediterranean) (Parks et al.,
2012). In the fossil record of European pines seed cones have been discovered that are morphologi-
cally similar to P. canariensis or intermediate between P. canariensis and P. roxburghii, including an
Upper Miocene cone from the Vienna Basin in Austria described as a variety of P. canariensis (Klaus,
1988). Presumably, there were several species related to P. canariensis in the mountains around the
Mediterranean, in W Asia and in the Himalayas and some of these became extinct while others
retreated to the Mediterranean coasts or, in this case, to the newly formed Canary Islands.

Pinus canariensis on an old lava flow in the

central part of Tenerife, Canary Islands. Photo
credit Aljos Farjon.
 europe and the mediterranean 151

Picea

The genus Picea is represented in Europe with only two species, one is widespread and the other
is a narrow endemic. A natural hybrid between P. abies in Europe and P. obovata in Asia (as here
defined) occurs in a transition zone between the two, mostly in NW Russia and is treated in the
chapter Mainland Asia and Japan. There are no species in the Mediterranean area. Successive
glacial periods during the Pleistocene (nearly) eliminated all species and caused one, P. omorika,
to retreat to a small area in the Balkans. The genus also retreated much further E than it once
had been, and in Central and SE Europe to the mountains. In recent times forestry plantations
have expanded the limits far beyond these historical limits and naturalizations are blurring the
natural boundaries of spruces in many parts of Europe.

Map EM-17. Picea abies [n=208,

m=200, h=26, 17??–2010]

Picea abies is a very wide-

spread species occurring in
large parts of Europe, most
extensively in Scandinavia
(Norway, Sweden and Fin-
land) and in NE Europe
from Estonia and Belarus
far into Russia. In Central
Europe its natural distribu-
tion is limited to mountains
(all lowland spruce forests
are planted). It occurs from
the French Hautes Alpes
to the E end of the Alps in
Austria, in the Jura and Vosges Mountains, in the Black Forest, the Bavarian Forest and Bohe-
mian Mountains, the Erzgebirge, the Carpathians and the mountains of the Balkans S to Albania,
Macedonia and Bulgaria. In most of these mountains this important timber tree has been planted
extensively and it is not always possible to decide from herbarium labels if the occurrence was
natural or not. Natural occurrences in Romania could not be mapped for lack of herbarium speci-
mens from that country in the many herbaria consulted. For the same reason the Balkans are
under-represented. Despite this, the map does show the natural limits of Picea abies in Central
Europe. Its westernmost location is in the Vosges Mountains in NE France where this species
crossed the Rhine. The pollen records from ancient peat and clay deposits show that P. abies
occurred further W in Europe in previous interglacial periods and retreated to refugia in Russia,
Romania, the Balkans and N Italy during glacial advances in the Pleistocene. On its return since
the last glacial maximum its westernmost limits were further E than before (Schmidt-Vogt, 1987).
Its inability to compete with more shade tolerant Abies alba and Fagus sylvatica has further lim-
ited its natural expansion into W Europe. When planted further W, e.g. in the Belgian Ardennes, it
will invade areas with shallow acid soil and high rainfall, such as the blanket bogs and heathlands
of the Hautes Fagnes, and if left uncut form natural spruce forests.
152 europe and the mediterranean

Forest with Picea abies in the Sumava

Mountains, Czech Republic. This forest,
mostly undisturbed in the Boubin Forest
Reserve, has stands of pure spruce (Picea
abies) as here, and mixed stands of Abies
alba and Fagus sylvatica elsewhere. The
spruces occupy the sites with shallow
soils and poor drainage. Photo credit
Aljos Farjon.

Map EM-18. Picea omorika [n=18, m=18, h=3, 1875–2010]

In contrast with Picea abies, P. omorika has a very lim-

ited distribution. It occurs in a small area of Bosnia
and Herzegovina (Republika Srpska) on either side of
the Drina River and a short distance across the bor-
der into Serbia along the Tara River. Its main distribu-
tion is in the large bend of the middle Drina River in
the Tara Mountains, between Visegrad and Perucac;
other scattered populations occur to the NW around
Brioznik. Its southernmost locality is at Stajkovac and
its northernmost locality is at Viogor. In all of the over
fifty known localities it forms small and isolated sub-
populations, sometimes pure but often mixed with Picea abies or Abies alba and less commonly
with Pinus nigra or P. sylvestris. It grows mostly on limestone but sometimes on serpentine at
altitudes between 300–1700 m. In the Pliocene and early Pleistocene, Picea omorika (or its very
close relative P. omoricoides Weber) was widespread in Europe, with fossils known from e.g. the
Erzgebirge and Lüneburg in N Germany. Now protected, partly in the Tara National Park, it was
logged in the past and replaced by planted Norway Spruce (Picea abies) with which it does not
compete successfully in many localities. This species is listed as Endangered (EN) on the IUCN
Red List. It is widely planted as an ornamental tree for its graceful conical shape.

Larix

Only a single species of Larix is native to Europe, with three varieties that in Eastern Europe tend
to be recognized as species. The genus is limited to Central and Eastern European mountains.

Map EM-19. Larix decidua var. decidua

(black) [n=47, m=46, h=11, 1873–2009];
L. decidua var. carpatica (red) [n=20, m=19,
h=6, 1872–1990]; L. decidua var. polonica
(yellow) [n=7, m=7, h=4, 1908–2010]

Larix decidua has a limited distri-

bution compared to other common
European conifers and is restricted
to the Alps, the Carpathians and a
small area in Poland. Larix decidua
 europe and the mediterranean 153

var. decidua occurs in the Alps. In the W Alps (France and Switzerland), larch forests are mainly
present in the central to southern parts, which have a subcontinental climate with cold and rela-
tively dry winters. In Austria and Italy Larix decidua var. decidua is more widespread, as the Alps
here are further from the Atlantic influence bringing in warmed air and heavy snowfall. Larix
decidua var. decidua often occurs with Pinus cembra (map EM-13 on p. 148) and has a similar
distribution. Both occur near the tree line, but this has been artificially lowered by centuries of
pasturing. Its highest limits are between 2200 m and 2500 m. Larix decidua var. carpatica has
a scattered distribution in the Carpathian Mountains from Upper Silesia (Sudeten Mountains)
in Poland to Romania. Its altitude range is well below that of larches in the Alps and it occurs
often together with Picea abies, Pinus cembra or P. sylvestris. Larix decidua var. polonica has a
very limited distribution on the headwaters of the Wisła River in Poland, at altitudes between
150–350 m. It occurs scattered in pine-birch forest or oak-birch forest and is in many places rare.
The geographical extent of this variety is uncertain and could be greater than presently known,
but records from the N slopes of the Tatra Mountains are better placed with var. carpatica. This
lowland form of the European Larch is probably a relict of colder times and is likely to be replaced
by those trees that now surround its last, often old individuals unless measures are taken to
stimulate regeneration. This variety is listed as Vulnerable (VU) on the IUCN Red List.

Abies

There are six species of Abies in Europe and the Mediterranean (excluding Turkey) including
one nothospecies, A. ×borisii-regis. Only one species, A. alba, is widespread, the others are more
limited in their distribution or even confined to a single locality. The most limited species occur
in the Mediterranean and those with somewhat wider distribution in the Balkans. It appears from
the fossil record that all species may have had much wider distributions and that there were more
species (see e.g. Liu, 1971 pp. 61–62), often found in places where they no longer occur naturally, e.g.
in the Miocene/Pliocene flora of the Massif Central in France (Legrand, 2010). North of the Alps
and Pyrenees only Abies alba survived, or returned from refugia after the last glacial maximum
10,000 years ago. The Mediterranean species of Abies are most likely relicts of a more diverse
conifer flora in Europe which started to become depleted by the ice ages of the Pleistocene.

Map EM-20. Abies alba [n=127, m=125,

h=20, 1842–2010]

Abies alba is widespread in Cen-

tral Europe including the Alps, in
the Pyrenees, the Carpathians, the
Balkans, Italy and Corsica. The
southernmost populations are in
Calabria, S Italy; the northernmost
in the lowlands of Poland around
51° N. In the Balkans introgression
with A. cephalonica during repeated advances of A. alba from the N in the Pleistocene has caused
its boundary to be unclear (the putative hybrid is here treated as A. ×borisii-regis, see map EM-21
on p. 154). Two localities in N Greece and W Bulgaria apparently have populations of A. alba
within or near the range of the nothospecies. In Italy, it is rare in most of the Appenines and
only more common or widespread in the N of that range (not shown on the map: no data) and
in Calabria. The species is rare in the Central Alps, where a drier and colder climate prevails.
Abies alba has been widely planted as a forestry tree and some published distribution maps of
the species distinguish poorly between natural and artificial occurrences. A more reliable map is
published in Hegi’s Flora von Mitteleuropa Vol. 1, part 2: 40 (Zoller, 1981). This map shows outlying
154 europe and the mediterranean

occurrences in the Massif Central and in Normandy (France) but it seems doubtful that these are
natural populations, especially the location in Normandy, as they are situated well within the
Atlantic climate zone where at lower altitudes than ca. 700 m beech (Fagus sylvatica) would pre-
vail (Ellenberg, 1988). Abies alba is a conifer of colline to montane altitudes (from 300 m in Poland
to 2000+ m in the Pyrenees and S Alps, partly as krummholz) where precipitation is abundant.
It forms pure forests or mixed forests with Picea abies and Fagus sylvatica. In some mountains of
Central Europe, e.g. the Harz Mountains in Germany, no natural populations occur despite suit-
able habitat, but it has been planted there, too.

Abies alba with Picea abies (in distance) and Fagus sylvatica
in the Sumava Mountains, Boubin Forest Reserve, Czech
Republic. It is early April before the beeches have come into
leaf. In this strict forest reserve no trees have been cut since
more than 150 years in order to enable the study of natural
succession processes. Photo credit Aljos Farjon.

Map EM-21. Abies ×borisii-regis [n=31, m=31,

h=7, 1891–2006]

Abies ×borisii-regis is a putative natural

hybrid between A. alba and A. cephalon-
ica, a stabilized nothospecies that breeds
true and forms natural populations. Its
distribution is situated between the pre-
sumed parent species, in S Albania, N and
NE Greece and SW Bulgaria (Rhodope
Mountains). Abies cephalonica occurs in
S Greece (map EM-22 on p. 155). During
periods of glacial expansion in the Pleistocene forests in Europe were forced S and E into refugia,
some of which were situated in the Balkans. During these prolonged cold phases A. alba came
into contact with A. cephalonica and hybridization occurred (Mitsopoulos & Panetsos, 1987). Its
habitat and place in the forest vegetation is very similar to A. alba, occurring in beech or beech-
chestnut forest at lower elevations (from 250 m upwards) and in conifer forest with Picea abies
or in pure stands up to 1800 m a.s.l. As the Balkan Peninsula is more diverse in tree species than
Europe N of the Alps, both angiosperm dominated and conifer dominated forests often have
other species besides the co-dominants. Yet it seems from the distribution maps based on her-
barium specimens that away from local contact zones between the nothospecies and its putative
parent species there is only A. ×borisii-regis indicating stabilized genetics in the firs in this region.
 europe and the mediterranean 155

On the other hand, some taxonomic bias may be present because herbarium specimens are rarely
‘named’ as intermediate between one and another taxon.

Map EM-22. Abies cephalonica [n=45, m=45, h=8,

1852–2008]

Abies cephalonica is distributed in S Greece,

from the southern Pindos Mountains in the N
(where it meets A. ×borisii-regis and perhaps
is no longer ‘pure’) through Sterea Hellas and
the Peloponnisos to the Taygetus Mountains in
the S. It also occurs on the island of Cephalonia
(Kefallonia) in the W and on Euboea (Evvoia) in the E. Much of its distribution is on limestone,
but towards its N limits siliceous rock also occurs with this species, which is more a montane
fir than A. ×borisii-regis and A. alba to the N, occurring between (600–)800–2000(–2100) m.
The climate is Mediterranean with most of the precipitation occurrring in winter. Presumably
A. cephalonica once had a greater distribution extending further N, where now A. ×borisii-regis
occurs. When summers become drier in a warming climate, A. cephalonica would seem to have
an advantage, but it is difficult to establish if it is already expanding. Forestry plantings of A. alba
and A. ×borisii-regis in mountains where A. cephalonica occurs will reverse such expansion in
favour of the hybrid trees.

Map EM-23. Abies nebrodensis [n=4, m=4, h=3, 1914–2000]

Abies nebrodensis is the rarest of the European/Mediterranean

firs. It was thought in the past to have been reduced to a
single tree in a village, but later a very small population was
found among clumps of oak (Quercus petraea) and beech
(Fagus sylvatica) on partly denuded limestone slopes in the
Madonie Mountains of N Sicily, Italy. Presumably, the tree in
the village of Polizzi Generosa, larger than any of the 29 individuals known in 2006 to still occur
on Monte Scallone on a slope above the Vallone Madonna degli Angeli, is a remnant of a more
extensive mixed forest in which A. nebrodensis occurred. The forest was cut long ago, after which
browsing goats prevented regeneration and erosion of the slopes made establishement of new
trees very difficult. This is now being attempted by outplantings from large stocks of seedling
trees (and cuttings) established in nurseries in various parts of Europe as well as in Sicily. This
species, presumably derived from a Late Pleistocene refugium of A. alba and closely related to it,
is listed as Critically Endangered (CR) on the IUCN Red List.

Map EM-24. Abies pinsapo var. pinsapo

(black) [n=21, m=21, h=7, 1837–2007];
A. pinsapo var. marocana (red) [n=17,
m=17, h=4, 1955–2006]

Abies pinsapo has a limited distri-

bution in S Spain and N Morocco,
on either side of the Strait of
Gibraltar in mountains that would
have been connected before the
break-through of the Atlantic
waters into the Mediterranean
156 europe and the mediterranean

Basin ca. 5.33 Ma. In Spain, A. pinsapo var. pinsapo occurs in the provinces of Cádiz and Malaga in
the following mountains: Sierra de Grazalema, Sierra de la Yunquera, Sierra de las Nieves, Sierra
Bermeja, Sierra Blanca and Sierra de Tolox. These are mountains of moderate height (highest
point 1919 m) and A. pinsapo var. pinsapo occurs there between 900 m and 1800 m a.s.l. It grows on
rocky soils derived from dolomitic limestone or serpentine. A number of threats, including over-
grazing and increased incidence of fires, caused this variety to be listed as Endangered (EN) on
the IUCN Red List. In Morocco, var. marocana has an even more limited distribution. It occurs in
the Western Rif Mountains, partly in Talassemtane National Park and on a few mountains outside
this reserve, most extensively on Jebel Tazaot. The Moroccan variety occurs at higher altitudes
than the Spanish variety, between 1400 m and 2100 m, and grows in more or less pure stands or
mixed with Cedrus atlantica. Despite its presence in a national park, threats to this variety are
similar to those in Spain and it is likewise listed as Endangered (EN) on the IUCN Red List.

Map EM-25. Abies numidica [n=7, m=7, h=6, 1861–1971]

Abies numidica occurs in NE Algeria in two localities in the Petite Kabylie Mountains near the
coast of the Mediterranean Sea: on the mountains Babor and Ta Babor. It grows in small forests
on the N-facing and E-facing slopes of these mountains on limestone, mostly between 1800 m and
2000 m, where Cedrus atlantica is the dominant conifer. In a few places A. numidica forms pure
stands, or grows with the shade tolerant common yew, Taxus baccata. Dry and hot summers alter-
nate with cool and wet winters, but it is evident that this species is limited to these two moun-
tains due to climatic constraints. Abies numidica is closely related to A. pinsapo and is probably
a relict of a much more widespread distribution of Abies in the Atlas Mountains bordering the
Mediterranean in W North Africa during the cold phases of the Pleistocene. The trees are legally
protected against cutting but forest fires and other destructive events have severely depleted the
two populations. It is listed as Critically Endangered (CR) on the IUCN Red List.

Cedrus

The genus Cedrus has three species in Farjon (2010a); traditionally four are recognized but there
may be as few as only two. Its distribution is highly disjunct, with populations in the Atlas Moun-
tains of Algeria and Morocco (C. atlantica), the Lebanon Mountains, Syria and S Turkey (C. libani),
Cyprus (C. libani var. brevifolia) and the Hindu Kush, Karakoram and Himalayas (C. deodara).
This distribution suggests contraction of a formerly more extensive and continuous range of pos-
sibly 1–2 species. The fossil record, although sparse as usual, provides evidence of occurrences
in France (Miocene), S Europe (Miocene, Pliocene), W Kazakhstan (Oligocene) and the Ahaggar
Massif in the central Sahara (early Pleistocene) according to Legrand (2003) and Florin (1963).
A possible migration route lay across the Thetys Fold Belt, a series of mainly Neogene mountain
uplifts along the brink of a closing Thetys Sea, once extending from SE Asia to NW Africa, and
of which the Black Sea and Mediterranean Sea are remnants. Desertification in more recent mil-
lennia separated the 3–4 major population areas further. The distance between C. atlantica and
C. libani is ca. 1800 km and that between C. libani and C. deodara ca. 2700 km. Phylogenetic analy-
sis (Qiao et al., 2007) using molecular sequences of cpDNA and mitochondrial DNA from samples
 europe and the mediterranean 157

of four species has placed C. deodara as sister to the Mediterranean taxa. No age estimate was
given for the split between this species and the other three, but the divergence between North
African C. atlantica and C. libani and its variety brevifolia was estimated to have taken place
18–23 Ma, i.e. in the Early Miocene. These results and the fossil record indicate an Asian origin
of the genus and migration to Europe, thence to North Africa and subsequent vicariance caused
by climate change, making large intervening territory unsuitable for these trees. In this chapter
we present the maps for C. atlantica and C. libani; C. deodara is treated in the chapter Mainland
Asia and Japan.

Map EM-26. The distribution of the genus Cedrus.

Map EM-27. Cedrus atlantica [n=41,

m=41, h=12, 1853–2006]

Cedrus atlantica occurs in the

central Haut Atlas, the Moyen
Atlas and the Rif Mountains of
Morocco and in the Atlas Tellien
of Algeria. Between these two
areas the species is absent along
a distance of ca. 700 km where the Atlas Mountains are much lower. More enigmatic is that
C. atlantica appears to be absent in the high W parts of the Moroccan Haut Atlas. The Atlas cedar
occurs at elevations between 1370 m and 2500 m a.s.l., especially on N and NW exposed slopes
receiving 1000 to 2000 mm precipitation annually, mostly during the winter. At the high ridges
much snow accumulates. The summers are warm and dry. Cedrus atlantica is often the dominant
conifer and forms pure stands or is associated with e.g. oaks, Populus tremula and Taxus baccata.
Heavy logging in the decades between 1940 and 1980 has reduced many stands of the Atlas cedar;
the decline may be as high as 75% in Morocco alone. This species was recently re-assessed as
Endangered (EN) for the IUCN Red List.

Map EM-28. Cedrus libani var. libani

(black) [n=57, m=55, h=10, 1853–2009];
C. libani var. brevifolia (red) [n=19,
m=18, h=2, 1879–2003]

Cedrus libani, the ‘classical’ Cedar

of Lebanon, occurs in mountains
bordering the E end of the Medi-
terranean Sea. It is native to Leba-
non, Syria, Turkey and Cyprus. In
Lebanon it occurs in the highest
parts of Jebel Liban (Mountains of
Lebanon) in isolated small forests or stands from Jabal el Barouk in the S to Mt. Akar in the N. In
Syria this species is limited to a few localities in Al Ladhiqiyah in the mountains E of Al Haffah.
158 europe and the mediterranean

The most extensive part of the range of C. libani var. libani is in the mountains of S Anatolia,
Turkey. Here it is present from the Nur Dağları (Amanus Mts. in classical texts) and Ahir Dagh E
of the city of Kahramanmaras in the E to Baba Dagh in Mugla in the W. An isolated population
occurs in N Turkey not far from the Black Sea coast in Tokat in the Canik Dağları N of Erbaa.
This northernmost locality is of interest as it appears to indicate that this species may have had a
much wider distribution than at present in NE Turkey, including possibly connecting populations
between the E Mediterranean and the Black Sea. The altitude range of this species is substantial,
from 1100 m to 3000 m. Cedrus libani is an endangered species in Lebanon and Syria due to mil-
lennia of over-exploitation, leaving the mountains virtually deforested. In Turkey exploitation has
had a much less severe impact and extensive forests occur in the Toros Dağları (Taurus Moun-
tains). In Cyprus occur a few small cedar forests in the Troödos Mountains and these populations
are recognized as C. libani var. brevifolia (or traditionally at species rank). Mainly due to a limited
area of occupancy (AOO) these populations are at risk from catastrophic events such as forest
fires and therefore listed as Vulnerable (VU) on the IUCN Red List.

Cedrus libani var. libani on limestone in the

Taurus Mountains of southern Turkey. Photo
credit Aljos Farjon.

Cupressaceae

This family is present in the region with three genera, Cupressus with two species, Juniperus with
nine species and Tetraclinis with one species. For the most part these species occur in coun-
tries around the Mediterranean Sea, but Juniperus brevifolia and J. cedrus occur on islands in
the Atlantic Ocean and J. sabina and J. communis extend much farther across Eurasia. Juniperus
communis is notable as the most widespread species of conifer in the world with a circumboreal
distribution. We treat it here as it was first described by Linnaeus from Europe and because our
data allow more distribution detail there than in most other parts of the Northern Hemisphere
in which it occurs.

Cupressus

The genus Cupressus in Europe, SW Asia and North Africa is represented by taxa with adaptations
to in some cases extreme drought or at least dry and hot summers. Desertification has reduced
the distribution of Cupressus in the region, followed by human exploitation of the landscape
especially in the Middle East and the Mediterranean, so that the present natural distribution is
not only limited but highly fragmented.
 europe and the mediterranean 159

Map EM-29. Cupressus sempervirens [n=62, m=55, h=14, 17??–2009]

Cupressus sempervirens occurs in the Mediterranean and in parts of SW Asia. It has been widely
cultivated as an ornamental tree in the region, almost invariably in its columnar fastigiated form
known as Italian Cypress, or selections with similar crown shapes. Although that branching habit
occasionally occurs in natural populations, from which it has undoubtedly been selected for plant-
ing long ago, the natural trees normally have spreading branches. Whether these were formerly
more widespread is not known, but at present they do not occur naturally in Europe except on
Crete, Karpathos, Rhodos, Kos, Kalimnos, Samos and perhaps a few other islands in the Dodeca-
nese Islands (Browicz & Zielinski, 1982). All other occurrences in Europe are planted or come from
planted trees of the columnar or narrowly pyranidal form and are excluded from the map. Cupres-
sus sempervirens is quite widespread in S Turkey, from Samsun Dagi in the W to Gavur Dağları
(Amanus Mts.) in the E, in the coastal mountains of Syria, in Lebanon, N Israel (but probably
mostly introduced), Jordan (two localities, not mapped) and on Cyprus. It occurs in a few localities
in North Africa, one in Tunisia (Maktar, perhaps not of natural origin) and three in Lybia, with
the more extensive stands in Al Jabal al Akhdar (Cyrenaica). Probably the most extensive forests
of C. sempervirens occur in Antalya, S Turkey. In Iran there are natural populations in the Elburz
Mountains around the S coast of the Caspian Sea, but also, very disjunct, 3–4 localities in S Iran
(two shown on map) which may be relicts of more extensive forest before desertification of much
of the region. Some of the trees there are very large and old, one with a trunk circumference of
12 m. As far as known, no natural stands are found in the Transcaucasus or in the Kurdistan region
where Iran, Iraq and Turkey share borders. Given a long history of human occupation it cannot be
ruled out that this species once occurred there but was exploited to extinction.

Map EM-30. Cupressus dupreziana var. dup­

re­ziana (black) [n=6, m=6, h=4, 1925–1985];
C. dupreziana var. atlantica (red) [n=21,
m=21, h=7, 1923–1997]

These two very disjunct and highly rel-

ict taxa have commonly been recog-
nized as species, but they are in fact
extremely similar and could even be
considered as subspecies or varieties of
C. sempervirens. Cupressus dupreziana
var. dupreziana occurs in the W-central
part of the Sahara in SE Algeria, on the
Tassili Plateau (Tassili n’Ajjer) N and NW
of the oasis of Djanet. The Tassili escarpment is the upended edge of a vast sandstone plateau
and is dissected by numerous canyons. Up on the plateau a maze of rock towers and sandstone
bluffs interspersed with sandy wadis and stony plains support an extremely sparse vegetation.
160 europe and the mediterranean

The cypresses grow in the wadis or in depressions where there is water underground. The most
recent survey (Abdoun & Beddiaf, 2002) counted 233 living trees and several seedlings and young
plants, indicating sporadic regeneration probably linked with rare episodes of rain fall. The alti-
tude at which these trees grow, some over 2000 years old, is 1700–1950 m. There is evidence, e.g.
in the famous prehistoric rock art which made the Tassili n’Ajjer a World Heritage site, of a wetter
climate in prehistoric times which enabled animals of the African savannah to exist here, fol-
lowed by cattle herding when people moved in. Presumably the high plateau was then forested;
the cypresses survive on the ground water which gets blocked from running off by rock ledges.
Cupressus dupreziana var. atlantica occurs in S Morocco in the Oued-n’Fis valley and adjacent
localities in the W end of the Haut Atlas. Here the vegetation, although adapted to dry conditions,
is more abundant than in the Sahara and probably not dissimilar to that which accompanied var.
dupreziana several thousand years ago. Common associates of var. atlantica are Juniperus phoe-
nicea, Tetraclinis articulata and Pistacia lentiscus, but the present situation is woodland degraded
by overgrazing and browsing from goats, causing loss of vegetation cover and erosion. Both variet-
ies of C. dupreziana are listed as Critically Endangered (CR) but the species is Endangered (EN)
on the IUCN Red List.

Cupressus dupreziana var. dupreziana

in a wadi on the Tassili Plateau, NE of
Djanet, Sahara, Algeria. Photo credit
Aljos Farjon.

Juniperus

Two species of Juniperus have distributions that extend far beyond the region discussed in
this chapter and these are mapped and discussed here first; the remainder has an Atlantic-
Mediterranean distribution. Juniperus, unlike its close relative Cupressus, is dispersed by birds
which may explain the wide distribution of some species and is certainly responsible for the
occurrence on oceanic islands which never had a connection to other lands.

Map EM-31. Juniperus communis [n=749, m=695, h=65, 17??–2010]

 europe and the mediterranean 161

Map EM-32. Juniperus communis in Europe. J. communis var. communis (black); J. communis var.
saxatilis (red)

Juniperus communis has the widest distribution of all conifer species. It is the only species that
is circumpolar and extends across Eurasia and North America. A circumpolar map is given in
Hultén & Fries (1986) but we present the species here in two maps, one with the global distribu-
tion of the species and one for Europe, here expanded to include European Russia, the Caucasus
and Turkey. For the latter region our data are more complete than for the global range of the
species and we separate the two varieties that occur there.
Juniperus communis has a boreal-temperate distribution, reaching from the Arctic tundra zone
to the Mediterranean. In Europe it occurs within the Arctic Circle in Iceland, Scandinavia and
Russia, in North America it reaches similar N latitudes in Alaska and Canada. Its southernmost
locations are on the North African coast in Algeria and on the border between Mexico and the
USA. In Eurasia it extends from Iceland, Ireland and Portugal in the W to Kamchatka and Japan
in the E; it also spans the North American continent from the Pacific to the Atlantic coast. Four
varieties are recognized in Farjon (2010a) and these appear to be geographically more or less dis-
tinct. Several additional varieties are recognized by Adams (2011) based on chemical and molecu-
lar data. In part due to incomplete specimen data for mapping and because of this taxonomic
flux, we do not separate the varieties on the global map but briefly mention their distributions
below. Juniperus communis var. communis occurs in Europe with extensions in North Africa, the
Caucasus region and NW Iran. Its limits in Russia possibly extend as far E as the Ural Mountains
but records are very sparse and it becomes very rare with increasing continentality of climate.
Northwards it is also limited to S Scandinavia and becomes rare in Scotland. This variety is con-
current with a relatively mild winter climate. Juniperus communis var. saxatilis occurs in Eurasia
and in North America. In Eurasia it has a more northern distribution than var. communis, but also
occurs in high mountains including the Himalayas, Hindu Kush, Tian Shan and Altai Mountains,
and extends much further E as far as Kamchatka, Sakhalin Island and Japan (var. nipponica). In
North America it is mainly distributed in the NW from Alaska to Oregon, but has been found in
Labrador and on the ice free coasts of Greenland. Juniperus communis var. depressa is widespread
in North America from inside the Arctic Circle to the Mexican border, but absent from the Pacific
Northwest of the USA (where var. saxatilis is found) and the Great Plains. Finally, J. communis
var. megistocarpa has a limited distribution in maritime Canada, where it is known from the
W coast of Newfoundland, Madeleine Island (Quebec) and Sable and Saint Paul Islands (Nova
Scotia). All varieties (including the additional ones recognized by Adams) except var. communis
are prostrate or decumbent shrubs and they occur in various habitats ranging from arctic tundra
162 europe and the mediterranean

and alpine rocky meadows to peat bogs, bare rock surfaces smoothed by glaciers to limestone
barrens and coastal sand dunes.
Juniperus communis in Europe is mapped here in more detail as more data from herbaria are
available. Two varieties occur here, var. communis and var. saxatilis. They are separated primar-
ily by climatic factors but where they are sympatric edaphic factors also play a part, so that they
only rarely occur together in the same locality. Juniperus communis var. communis is an upright
shrub to a small tree; var. saxatilis a prostrate shrub. The northern limit of var. communis is at
65° N in Karelia, NW Russia, but further W the line follows S Finland, the Åland Islands, across
to Sweden and the coast of Norway to Møre & Romsdal. The increasing Atlantic influence causes
var. communis to be replaced by var. saxatilis in Shetland, the Faroes, Iceland, Scotland and the
W coast of Ireland. Juniperus communis var. communis requires warm summer days for its cones
to mature and these occur in lowlands of the more southern and/or continental parts of Europe.
Juniperus communis var. communis and var. saxatilis are therefore separated by altitude at more
southern latitudes, where the latter variety becomes a high montane to subalpine representative
of the species. Here var. saxatilis occurs e.g. in the Pyrenees, Alps, Carpathians, Corsica, the Bal-
kan Mountains and across N Turkey but is peculiarly absent in the Taurus Mountains of S Turkey.
In the lowlands J. communis var. communis occurs on poor sandy soils or on calcareous soils in
open vegetation types, e.g. heaths and chalk grasslands, and can maintain itself in open birch-
pine or birch-oak woodland, but eventually disappears when the forest canopy closes. As less
intensive traditional land uses are being replaced by a mosaic of intensive agriculture and man-
aged or unmanaged dense woodland, the common juniper becomes less common in many parts
of Europe.

Juniperus communis var. communis in

the Dwingelo Heath, Drenthe, Nether­
lands. Photo credit Aljos Farjon.

Map EM-33. Juniperus sabina var. sabina (black) [n=196, m=168, h=26, 17??–2009]; J. sabina var. arenaria
(red) [n=13, m=12, h=6, 1918–1998]; J. sabina var. davurica (yellow) [n=19, m=13, h=9, 17??–1993]
 europe and the mediterranean 163

Juniperus sabina has a very extensive longitidunal distribution from Morocco in the W to Pri-
moriye in the Russian Far East, a distance of more than 10,000 km. Its latitudinal distribution
is relatively limited, from around 32° N in Morocco to around 54° N in European Russia. The
distribution of J. sabina var. sabina is nearly as extensive but extends in the E not further than
Mongolia and Shaanxi Province in China. In the Gobi Desert appears var. arenaria, and in the far-
thest E in Nei Mongol (Inner Mongolia) and Heilongjiang Province in China and in Primoriye in
Russia occurs var. davurica (J. davurica in Chinese and Russian publications). In Europe, J. sabina
is a high montane to subalpine prostrate species occurring in Spain, France (Pyrenees and Alps),
the Alps of Switzerland, Italy and Austria, the Carpathian Mountains of Romania and Ukraine,
and mountains in the Balkan Peninsula. Plants identical with var. davurica have been found in
E Tirol, Austria, but so far these are the only records and all other collections in the Conifer Data-
base made in Europe belong to var. sabina. J. sabina var. davurica is possibly a recurrent juvenile
leaf form (Farjon, 2005a). The nominal variety (sabina) is also a montane species in N Turkey, the
Crimea, the Caucasus, N Iran, Central Asia and the Altai Mountains. However, from Volgograd
Province in Russia eastwards it occurs here and there in the steppes, often on sand dunes near
rivers and on calcareous hills; these are the northernmost localities. As is the case with J. commu-
nis var. saxatilis, J. sabina has not been found in the Taurus Mountains of S Turkey. Other ‘gaps’
where we would expect this species are the W Carpathians, Calabria in Italy, and the mountain
ranges in Uzbekistan and W Kyrgyzstan. There is the possibility of missing records in some of
these regions, but at least S Turkey seems genuinely devoid of this taxon. An explanation for this
absence cannot readily be given; it is possibly an accident of dispersal history.

Map EM-34. Juniperus oxycedrus ssp. oxycedrus (black) [n=196, m=185, h=22, 17??–2008]; J. oxycedrus ssp.
macrocarpa (red) [n=67, m=55, h=19, 16??–2009]; J. oxycedrus ssp. badia (yellow) [n=17, m=17, h=5, 1853–
2002]; J. oxycedrus ssp. transtagana (blue) [n=15, m=13, h=11, 1838–2002]

Juniperus oxycedrus is a variable species; here we present a map with four commonly recognized
subspecies. The species is distributed in a ‘greater’ Mediterranean/Black Sea region, extending
from Portugal and Morocco to the Kurdistan region of Iran, Irak and Turkey. The most wide-
spread subspecies is J. oxycedrus ssp. oxycedrus, which occurs on the coast as well as inland and
from near sea level to 2200 m. It is common in rocky terrain with maquis or open sclerophyllous
woods dominated by Quercus ilex, Pistacia lentiscus and Carpinus betulus and in clearings within
Pinus brutia or P. halepensis forest, but also appears in montane and wetter forest with Cedrus
atlantica or C. libani, Pinus nigra, Juniperus foetidissima, and J. excelsa. At higher altitudes and
inland this subspecies is subject to winter frost, yet it has not spread northwards beyond ca. 45° N
even where it is not barred from this by high mountains, e.g. in S France. Juniperus oxycedrus
ssp. macrocarpa occurs along the coasts of the Mediterranean and Atlantic Ocean from S Spain
to S Turkey. Unlike ssp. oxycedrus it is never far from the sea and its highest recorded location is
75 m; more commonly it grows on beaches and sand dunes between 1–10 m. Very few herbarium
collections are known from the North African coast, so it is much more common on the European
164 europe and the mediterranean

side of the Mediterranean Sea, but even here it is very scattered on some coasts, such as in Spain
and on the Adriatic coasts. The two other subspecies are more restricted; J. oxycedrus ssp. badia in
Portugal, Spain and Algeria and J. oxycedrus ssp. transtagana in a small area from central Portugal
to S Spain, mostly on or near the coast of Portugal in Setúbal. While ssp. badia occurs in inland
Spain at altitudes up to 1300 m, ssp. transtagana is mostly confined to low, sandy areas where it
occurs in open pine woods or coastal grass and scrub.

Map EM-35. Juniperus phoenicea ssp. phoenicea (black) [n=191, m=172, h=25, 1700–2009]; J. phoenicea ssp.
turbinata (red) [n=38, m=38, h=15, 1845–2006]

Juniperus phoenicea is a Mediterranean species with a distribution which extends westwards to

the Canary Islands and Madeira and eastwards to the Arabian coast of the Red Sea. In Europe
this species occurs from Portugal in the W to Crete and the Dodecanese Islands in the E; it is
also present in W and S Turkey, on Cyprus, in N Syria, Jordan, Israel and the Sinai Peninsula of
Egypt (the only conifer native in that country). Similar to J. oxycedrus ssp. oxycedrus, the more
widespread nominal subspecies J. phoenicea ssp. phoenicea occurs on coasts and islands as well
as inland, while J. phoenicea ssp. turbinata is a taxon restricted to sandy beaches and coastal
dunes. The occurrence of J. phoenicea on the islands in the Atlantic Ocean is undoubtedly due
to dispersal of the seeds, packaged in succulent red cones, by migrating birds. Like J. oxycedrus
ssp. macrocarpa, J. phoenicea ssp. turbinata is rare on the North African Coast, where sandy sea
shores are much more extensive than in Europe. There may be some collection bias, but the
overall pattern seems genuine and environmental factors related to the nearby Sahara Desert are
possibly responsible. Where the desert is close to the sea, the species is absent; in Saudi Arabia
it is limited to the high escarpments of the Hijaz and Asir which receive some moderate winter
rain, unlike the Nubian Desert on the W side of the Red Sea. In Morocco and Algeria J. phoenicea
ssp. phoenicea occurs in the Atlas Mountains from ca. 1000 m to 2150 m a.s.l.; in the Asir Range
of Saudi Arabia even to 2400 m, together with the African tree juniper J. procera (map AF-5 on
p. 462). Juniperus phoenicea is a shrub or small tree of maquis or carrique or sclerophyll open
woodland; in the higher mountains of the Atlas Range also in conifer forest with Cedrus atlantica
and in Spain also in pine and oak forests.
 europe and the mediterranean 165

Map EM-36. Juniperus thurifera [n=63,

m=55, h=21, 1800–1997]

Juniperus thurifera occurs in SW

Europe and NW North Africa. This
species is most common and wide-
spread in Spain and Morocco and
more localized and rare in France
and Algeria. In Spain its distribu-
tion is in the N and E of the country,
but it avoids the wetter Cordillera
Cantabrica and Pyrenees as well
as the dry plains and plateaus of
Extremadura and La Mancha. Why
it is absent in the mountains of S
Spain is less clear, because it is
widespread further S in the Atlas
Mountains of Morocco. In Algeria
this species only occurs in the E part of the Atlas Range on two mountains in Aurès and to the
N in the Forêt de Lamblèche near Constantine. In France, most populations are in the Hautes
Alpes and Isère and very few on the N side of the Pyrenees. Juniperus thurifera is a large shrub or
tree to 20 m tall, occurring in semi-arid woodland with Quercus ilex and in the Atlas Mountains
also with Cedrus atlantica. In Spain it is mostly found on limestone, but in Morocco also on other
types of rock. Its altitude range is extensive, from 300 m to 3300 m (the highest occurrences in
the Atlas Mountains of Morocco). In France, this species is considered threatened with extinc-
tion, mainly due to changes in land use which cause forest succession to move in a direction of
taller, closed-canopy forest. If this is correct, then the wider occurrence of J. thurifera in the past
was due to human interference in the landscape and its vegetation, mainly through cutting and
the grazing of animals. The reverse is happening in Morocco, where too intensive cutting and
grazing threaten to suppress regeneration and in some areas only (very) old trees remain in a
half-denuded landscape.

Map EM-37. Juniperus foetidissima [n=100, m=85, h=17, 16??–2009]

Juniperus foetidissima occurs in the S Balkans in Europe, in Anatolia in Turkey, around the eastern
Black Sea from the Crimea to Georgia and NE Turkey, in the Lesser Caucasus and in Lebanon
and Cyprus. A few outlying localities are in the Elburz Mountains of Iran and Turkmenistan. In
Turkey this species is common in the W and becomes rare further E but is again common in
166 europe and the mediterranean

Georgia, Armenia and Azerbaijan. This species is the largest juniper in the region and some trees
in SW Turkey exceed 40 m in height and 4.5 m diam. and one such tree at Kadıncık Deresi is
estimated to be over 2000 years old (Farjon, 2005a: 77). Juniperus foetidissima is an upland spe-
cies usually occurring at altitudes between 600 m and 2000 m and occurs in open maquis or oak
scrubland as well as in woodland and conifer forest.

Juniperus foetidissima among cedars in the moun-

tains Bei Dağları, SW Turkey. Photo credit Aljos
Farjon.

Map EM-38. Juniperus drupacea [n=44,

m=36, h=11, 1787–2007]

Juniperus drupacea is a phyloge-

netically distinct species (Adams,
2011) with a disjunct distribution in
the eastern parts of the Mediterra-
nean Basin. It occurs in the Par-
non Mountains in Peloponnisos,
Greece, in S Turkey especially in
the Taurus Mountains, in W Syria
(mostly in the Jabal an Nusay-
riyah), in Lebanon and at one
locality in Israel near Nazareth (Galilea). The distance between the Greek population and the
nearest occurrence in Turkey is ca. 800 km and it is likely that J. drupacea has occurred in areas
in between these locations. It is also notably absent from islands such as Crete and Cyprus where
remnants of ancient E Mediterranean coniferous forests characterized by Cupressus sempervirens,
J. foetidissima, J. excelsa, Abies cephalonica, A. cilicica, Pinus nigra, P. brutia and Cedrus libani still
occur. This is its main habitat in the Taurus Mountains of S Turkey, where such forests are more
 europe and the mediterranean 167

or less intact and include nearly all of these species. In Crete, the Peloponnisos and Cyprus, but
also in Syria and Lebanon, these forests are not only much reduced but also impoverished so that
several of these conifer species are absent. Juniperus drupacea can establish itself in more open,
shrubby vegetation which has replaced the forest and it has probably spread into this second-
ary vegetation from remaining forest remnants. Juniperus drupacea has exceptionally large seed
cones which contain three large and very hard, partly fused seeds. The fused bract-scales have
high sugar content. These characters suggest dispersal by mammals rather than birds and con-
sequently a more limited radius which could explain the absence of this species on the Aegean
and other islands.

Map EM-39. Juniperus brevifolia [n=31,

m=27, h=7, 1838–2010]

Juniperus brevifolia is endemic

to the Azores, the westernmost
group of islands in Macaronesia.
These volcanic islands are mostly
situated on the Azores microplate
on the E side of the Mid-Atlantic
Ridge; the two westernmost islands
are on the American plate W of
the ridge. The islands arose above sea level between 7 Ma (Santa Maria) and 300,000 years ago
(Pico); Santa Maria was submerged for a time and then re-emerged. Of the nine major islands,
seven have populations of J. brevifolia, these are from E to W São Miguel, Terceira, São Jorge, Pico,
Faial, Corvo and Flores. Juniperus brevifolia is related to J. oxycedrus and evolved on the islands
after seed dispersal by birds. The shortest distance at present between São Miguel and Portugal
is ca. 1350 km. Juniperus brevifolia occurs from near sea level to 1300 m on Ponta de Pico. The
prevailing SW winds (‘westerlies’) set the conditions for a moist but mild climate with abundant
rain and fog. Before humans arrived in the 16th century, this species was a constituent of the
Macaronesian evergreen rainforest of which only scattered remnants survive. The species is now
more commonly found as a shrub in secondary heathland and of larger trees only some stumps
remain. Ongoing habitat degradation causes this species to be listed as Vulnerable (VU) on the
IUCN Red List.

Map EM-40. Juniperus cedrus [n=34, m=30, h=10,

1838–1997]

Juniperus cedrus occurs on Madeira and in

the Canary Islands on Gran Canaria, Tenerife,
Gomera and La Palma. Madeira is ca. 450 km N
of Tenerife and is the largest island of a sepa-
rate archipelago. All these islands are of volca-
nic origin; the highest volcano is the still active
Pico del Teide on Tenerife at 3718 m which rises
inside the N rim of a giant caldeira and is snow-
capped for much of the year. Juniperus cedrus
has been a constituent tree 20–25 m tall in Mac-
aronesian lauraceous evergreen forest, mainly
on drier, rocky sites where this forest was less
well developed. This forest only remains intact
in a few places in deep canyons and the juniper
now mostly occurs in a stunted or shrubby form
168 europe and the mediterranean

on steep rock formations ranging in altitude from 450 m to 2400 m. Population sizes vary, the
smallest being the one on Gran Canaria where fewer than 10 mature trees are known to exist.
Since our dataset contains 15 herbarium collections made more than 100 years ago, there is the
possibility that some dots on the map represent populations that no longer exist. Apart from the
cutting of trees for its rot resistant wood, no longer practised, overgrazing by goats, some of them
feral, is the main problem and in some places the junipers only remain on the steepest rock faces.
Juniperus cedrus is listed as Endangered (EN) on the IUCN Red List.

Juniperus cedrus in the rocks of the

caldeira Cañada del Teide, Tenerife,
Canary Islands. Whereas in the past this
species was most commonly a tree in
the evergreen lauraceous forests of the
islands, with the virtual disappearance of
these forests due to human exploitation it
has now retreated to nearly inaccessible
rock slopes where even goats do not care
to climb. Photo credit Aljos Farjon.

Tetraclinis

Map EM-41. Tetraclinis articulata [n=83, m=78, h=13, 17??–2012]

The genus Tetraclinis has a single species, T. articulata, which is widespread in Morocco, N Algeria
and N Tunisia but occurs in only two locations in Europe: Sierra de Cartagena in Murcia, Spain
and on the island of Malta. In Morocco, the species occurs in the Atlas Mountains from just E and
N of Agadir to the border with Algeria S of Oujda and in N Morocco in the Rif Mountains as far N
as Djebel Dersa near Tetouan. The distribution in North Africa continues E through the Algerian
Atlas Mountains, but T. articulata becomes rare in the Monts des Nementcha in the easternmost
part of the Atlas Mountains. Fewer available herbarium collections may bias this distribution
pattern to some extent. Another gap between there and the population in Tunisia appears to be
real as all collections from that country are concentrated in the NE. Here it occurs in hills from
ca. 50 m to 200 m a.s.l. unlike in the Atlas Mountains where it can ascend to 1800 m (to 1300 m
on N-facing slopes). On Malta it is only known from a few localities with very few plants, some
 europe and the mediterranean 169

hidden in dolines or sink holes in the limestone barrens. It was thought to be almost extinct here
but recent surveys have discovered ‘new’ localities. In Spain it is also very restricted, with just
three localities known in the Sierra de Cartagena just E of Cartagena, all within a 20 km distance.
As the localities in Tunisia and Malta indicate, T. articulata is a conifer of warm, sunny terrain
and in the Atlas Mountains it occurs on the driest slopes. Rare in conifers, this species coppices
and survives with partly subterranean burrs or stools (an adaptation to fire) which humans have
exploited since prehistoric times. While the populations in Europe are threatened with extinc-
tion, the widespread abundance of T. articulata in North Africa places this species globally out-
side a threatened category on the IUCN Red List.

Taxaceae

The family Taxaceae is in this region represented by just one species, Taxus baccata. The genus
Taxus has the widest distribution of the genera in this family and is circumpolar, i.e. present in
North America as well as in Eurasia and North Africa. A distribution map of the genus Taxus is
presented in the chapter Global and trans-continental distributions.

Taxus

Map EM-42. Taxus baccata [n=265, m=261, h=26, 1831–2010]

Taxus baccata is widespread in Europe, occurring from W Norway to Sicily and from Portugal to
Romania. Beyond Europe it is present in the Azores and on Madeira in the Atlantic Ocean, in the
Atlas Mountains of North Africa, in Anatolia, the Caucasus and Lesser Caucasus and as far E as
the Elburz Mountains of N Iran. Its natural limits are primarily set by climate. In the N and E it
is limited by low temperatures; in Scandinavia T. baccata only leaves the coast in Denmark and
S Sweden and it is absent in European Russia. Whereas in N Europe it occurs in the lowlands, in
S Europe it retreats to mountains with cooler and wetter conditions, often in moist ravines. In
the lowlands of Central and NW Europe natural stands are often confined to chalk or limestone
or along small streams in loess or loam (Ellenberg, 1988). In recent times yews are spreading
widely into woodlands on more acidic sandy soils, coming from seeds dispersed by birds from the
ubiquitous plantings in churchyards, parks and gardens. We have endeavoured to exclude both
planted and naturalized yews from the database, but this is not always obvious from herbarium
specimen labels. In the NW and Central European lowlands Taxus baccata, while shade tolerant,
has been replaced by Fagus sylvatica and Carpinus betulus, shade tolerant angiosperms that grow
faster and taller, forcing T. baccata on to steep chalk slopes with a minimal soil development. It
was out-competed where these conditions did not exist; the present invasion of yews into utilized
170 europe and the mediterranean

but not intensely managed lowland woods will not last if these develop back into beech-domi-
nated forests. While yew wood was considered valuable, the toxic alkaloids in every part but the
red arils around the seeds are lethal to man and horse (deer and sheep are not affected) which has
led to early eradication from woods as well as planting in churchyards (death association). This
too has led to a shift in the natural distribution, which is therefore difficult to reconstruct.

Taxus baccata (darker green trees) in

mixed woodland (with Fagus sylvatica
on top of hill) on chalk, Mickleham
Downs, North Downs, England. Photo
credit Aljos Farjon.
Mainland Asia and Japan

Geography

The continent of Asia is the largest landmass in the world, with a total area of 46.65 million km²
as here circumscribed, including European Russia, Japan and smaller satellite islands such as Sri
Lanka, Hainan, Taiwan and Sakhalin. Europe is in terms of physical geography a part of this con-
tinent (Eurasia) but is treated in a separate chapter in this Atlas. The Malay Archipelago, usually
(partly) included in area calculations for the Asian continent but a distant archipelago, is also
treated in a separate chapter Malesia. Asia, like Europe, is thus a more or less artificial concept,
unlike the other five continents. Geographically it is more diverse than any of these due to its
vast extent across all climate zones and its complicated topography. Mainland Asia extends over
ca. 77° of latitude and ca. 195° of longitude or around 8600 km N to S and 9700 km E to W. In
the far N Russia borders on the Arctic Ocean and arctic desert and tundra merges to the S with
the greatest expanse of mostly coniferous taiga forest in the world. This region is further charac-
terized by great lowland rivers and generally low, old mountain ranges, both with a mostly S-N
orientation, but in European Russia the rivers run southwards. To the E Siberia is extended by the
great Kamchatka Peninsula, one of the most active volcanic regions in the world. Southwards, the
taiga merges to steppes in Kazakhstan and Mongolia and to high mountains such as the Caucasus,
Tien Shan and Altai, which have a mainly E-W orientation. Inland seas (Caspian, Aral, the latter
drying) and great lakes (Balkhash, Baikal) lie in the steppe zone or in the transition between taiga
and steppe (Baikal). Great deserts span almost the entire width of Asia at middle latitudes, from
the Arabian Desert to the Gobi Desert, with some of the driest and harshest deserts among them
(Rub al Khali, Takla Makan). Some of these deserts are mountainous (Zagros Mountains, Plateau
of Iran) or mountains surround the desert entirely (Takla Makan) and rivers running from them
do not drain to the ocean. In the S-central part of the continent rises the Tibetan Plateau, with an
area of 2.5 million km² the largest expanse of extremely high country in the world with an average
altitude of >4500 m. The Kunlun Range borders it in the N but its most remarkable edge is in the
S, formed by the Karakoram–Himalayas with 14 summits over 8000 m tall, culminating in 8848 m
at Mt. Everest (Chomolungma). These mountains are flanked in the W by the Hindu Kush and in
the E by a series of high mountain ranges trending NW-SE, interspersed by long rivers that have
their sources on the Tibetan Plateau. The subcontinent of India, forming an extension of Asia
into the Indian Ocean, lies to the S of these great mountains and its drift into the Asian Plate in
the early Palaeogene and subsequent pressures have warped them up to their great heights. The
subcontinent lies mostly in the tropics and is one of the most densely populated regions of the
world. The Deccan Traps, a volcanic plateau of great extent, the Western Ghats and the deltas of
the Indus and Ganges Rivers are its main topographical features. Eastern Asia has a complicated
topography. From the Manchurian Plain in the N to Hainan Island in the S, this part of China
reaches from boreal forest to tropical rainforest. Numerous mountain ranges, many trending N-S,
alternate with broad valleys of major rivers and plains such as the Red Basin and the Great Plain
172 mainland asia and japan

of China. In Sichuan/Yunnan the mountains rise to over 7000 m. This E half of China is also the
region with the largest human population in the world and, like India, it has been under culti-
vation for millennia, profoundly altering the landscape and the vegetation. Lastly, mainland SE
Asia (Indochina) is a broad peninsula extended by a narrow one, the Malay Peninsula, which
leads the continent to its southernmost point at Cape Piai at 1° 16’ N. Its E margin (Viet Nam) is
mountainous, as is much of the Malay Peninsula. The Mekong, originating on the Tibetan Plateau,
winds its long way through SE Asia forming the Mekong Delta in the S of Viet Nam. Major islands
border the Asian continent in the S and E and of these, the group forming Japan with Honshu the
central and largest island, is of special interest for conifer biogeography. Japan is mountainous
and largely volcanic, like Kamchatka lying in the ‘Ring of Fire’ around the Pacific Ocean. Strings
of smaller islands (Kuril Islands, Ryukyu Islands) of volcanic origin link Kamchatka, Japan and
Taiwan, the latter island mountainous but not volcanic.
Asia can be divided into regions in many ways but here we try to do so with the distribution of
conifers in mind. The diversity of conifers in China (including Taiwan) is such that it merits sepa-
rate treatment, even though its land boundaries are mostly not natural. Another obvious choice
is to treat the conifers of Japan separately, as these islands have a high level of conifer ende-
mism. The Himalayas, including the Hindu Kush and Karakoram Ranges, form a natural region
of extremely high mountains with many endemic or near endemic conifer species. Mainland
SE Asia (Indochina) merits separate treatment for the same reason, with mostly distinct conifer
distributions. Elsewhere, regionalisation becomes more diffuse for conifers and the regional divi-
sions made here are somewhat arbitrary for several species. Their outlines are, where necessary,
explained at the beginning of these chapter sections. The conifers in this chapter are mapped and
discussed within the following regions and in that order:
Western Asia, Central Asia and Siberia, Himalayas, China and Taiwan, Indochina, Mainland
Asia Far East, Japan.
Within these regions the taxa are presented in a linear sequence reflecting phylogenetic rela-
tionships, following Christenhusz et al. (2011). Statistics are given for China and Taiwan and for
Japan, the two regions with greatest conifer diversity.

Conifers in Mainland Asia and Japan

Families Genera Species Species + infra Countries with Area of occupancy Endemic taxa
conifers in km²
6/8 36/70 202/615 283/794 39/45 107,925/46,648,350 256
Numbers after ‘/’ are global or continent totals.

Mainland Asia and Japan exhibit high levels of diversity in conifers with six out of eight families,
just over half of all genera and nearly a third of the world’s species present. Conifers occur in the
wild in almost all countries, the exceptions are Kuwait, Bahrain, Qatar, the United Arab Emirates,
Sri Lanka and the Maldives. In many Asian species infra-specific taxa are recognized. This takes
the total of taxa below the rank of genus to 283 or 35.6%, more than a third of the world’s total.
Of these 256 or 90% are endemic to the region. This is to be expected for such a vast area, with
only a few species overlapping with Europe, one species also in Africa ( Juniperus procera) and
some more jointly in Malesia. Conifer species are very unevenly distributed across the continent
(map MAJ-1 on p. 173), with regions of high concentration contrasting with those poor in species
or even devoid of conifers. The map, it must be emphasized once more, shows the distribution
of species, not of trees; if it did the latter the boreal north of the continent would be filled with
dots. There are not enough herbarium specimen data to fill Siberia more densely to reflect the
taiga ­forest, which is poor in species numbers. At lower latitudes (from 45° N) the map is more
accurate. Conifer species are concentrated in Anatolia, W Syria and Lebanon, in the Caucasus and
 mainland asia and japan 173

Map MAJ-1. Distribution of conifer species in mainland Asia and Japan.

Trans-Caucasus, the Tien Shan and Altai, the Hindu Kush-Karakoram-Himalaya chain, in many
parts of mainland China plus Hainan and Taiwan, in Viet Nam, Japan and in Primoriye in the Rus-
sian Far East. These are predominantly mountain ranges or mountainous areas; it is evident that
conifer species are concentrated in montane environments, especially in the lower latitudes.
Just as conspicuous as the concentrations of conifers are the absences, perhaps even more so
where this absence is not expected. The deserts of Asia are for the most part too extreme and
conifers are absent in the Syrian and Arabian deserts, on the Plateau of Iran, the Kara Kum and
Kyzyl Kum E of the Caspian Sea, the deserts in S Afghanistan and Baluchistan (Pakistan), the Thar
Desert of India and Pakistan, the Takla Makan Desert in Xinjiang Province, China and in most
of the Gobi Desert. Other large areas without conifers are the Kirghiz Steppe covering most of
Kazakhstan and steppes around the northern end of the Caspian Sea and much of the steppes in
Mongolia and Inner Mongolia, although in the latter two a thin scattering of Juniperus sabina var.
arenaria is present. A striking absence is seen on the Tibetan Plateau, with no conifers occurring N
of the Yarlung Zangbo (Brahmaputra) River drainage. They continue to be absent in the Kunlung
Shan to the N of the plateau. A combination of extreme altitude and low annual precipitation and
temperatures are the causes, even though some conifers occur at similar or higher altitudes to the
S in the Himalayas and SW China. More remarkable is the near absence of conifers in the Indian
subcontinent. If this subcontinent is defined as the land between the Indus River in the W, the
Himalayas in the N and the delta of the Ganges in the E, the only area with a naturally occurring
conifer is in a limited area of the Western Ghats (Kerala and Tamil Nadu States, from ca. 11° 24’
N to ca. 9° 23’ N). The species there is Nageia wallichiana, widespread in SE Asia and most likely
the result of long distance dispersal in relatively recent time. Conifers are abundant in the fossil
record of India from the Lower Jurassic until the deposition of the Deccan Traps 68–60 Ma. They
have been assigned to families still known from the Southern Hemisphere including Taxaceae,
which is now mostly northern but with one living representative in New Caledonia. The youngest
fossils of conifers were found in the Deccan Intertrappean Beds, sediments formed during inter-
vals of these latest Cretaceous to early Palaeocene massive outpourings of flood basalts (Sitholey,
1963). No later conifers are known from the fossil record of India (Florin, 1963) and it is likely that
they became extinct during that time. The Indian subcontinent moved across a hotspot now in
the Indian Ocean when it was entirely in tropical latitudes and without substantial mountains.
When volcanism subsided, angiosperm rainforest must have dominated the landscape as it does
now in the Amazon and Congo Basins, leaving no ecological space for conifers.
The two maps MAJ-2–3, presenting diversity at the genus and species levels, show that much of
mainland Asia has low values and that diversity is mainly concentrated in China. Further concen-
trations are situated on two islands: Honshu in Japan and Taiwan. Moderately diverse areas are
174 mainland asia and japan

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Map MAJ-2. Map of Mainland Asia and Japan showing numbers of conifer genera per one degree cell.

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Map MAJ-3. Map of Mainland Asia and Japan showing numbers of conifer species + infraspecific taxa per
one degree cell.

Map MAJ-4. Rarity values per one degree cell. Cell score is the total of the rarity calculations for each spe-
cies in the cell. The rarity calculation is 1 divided by the total of cells occupied by a species. Thus, a species
that occurs in one cell scores 1 while a species that occurs in 10 cells scores only 0.1.
 mainland asia and japan 175

found in Turkey and the E Himalayas. The same pattern is found in map MAJ-4 which shows areas
of local endemism (species with small extent of occurrence or EOO). Within mainland China the
largest area with both high diversity and endemism is situated in the mountainous region that
extends from S Gansu through W Sichuan and NW Yunnan to the SE corner of Xizang [Tibet].
Although diminished, this trend extends westward along the Himalaya as far as Nepal. Wheras
Taiwan does not score very high in the species diversity map it is a hot spot for local endemism.
Endemism is high in the same areas of Honshu in Japan that show high diversity of species. The
correlation between diversity and endemism, while evident, is not present in all areas at all lev-
els. In particular, local endemics in E China and Viet Nam often occur in areas with moderate or
even low diversity. Although local endemics add to the diversity, they are not the primary drivers
of high diversity; most species in diverse areas have larger ranges. The narrow endemics, when
investigated individually, often turn out to be relicts with a wide distribution in the geological
past. It appears therefore, that high diversity of conifers in Mainland Asia and Japan is primarily
the result of relatively recent speciation. Phylogenetic analysis incorporating molecular dating
techniques has supported this (Leslie et al., 2012). The distribution of high diversity and to some
extent endemism is largely sympatric with areas of active orogeny and volcanism.
176 mainland asia and japan

Western Asia

Geography

Western Asia in this chapter includes the region so defined by the United Nations (http://
en.wikipedia.org/wiki/Western_Asia) plus Iran and the Russian part of the Caucasus. It com-
prises Turkey in Asia, the Caucasus, the countries on the E Mediterranean seabord, the Arabian
Peninsula and Iran. Large parts of this region are extreme deserts in which no conifers occur.
Turkey in Asia (Anatolia) is on three sides bounded by seas, with narrow sea straits separating
it from SE Europe. It is largely a high plateau surrounded by mountain ranges which are most
extensive in the N (Pontus Mts.) and S (Taurus Mts.) with Mt. Ararat in NE Turkey rising to
5137 m. The climate ranges from Mediterranean in the W and along the coasts of the Black Sea
and Mediterranean Sea to continental in the interior, with deminishing precipitation from W to
E. The Caucasus is a high mountain system culminating in Mt. Elbrus (5642 m), with the main
range (Greater Caucasus) separated from a smaller range (Lesser Caucasus) to the S. The latter
connects in Azerbaijan and N Iran to the Alborz (Elburz) Mountains around the S shore of the
Caspian Sea, in which Mt. Damavand (5610 m) is the highest volcano in Asia. The slopes facing the
Caspian Sea receive abundant precipitation and are forested; on the S side the mountains merge
with the semi-arid Qazvin-Tehran Plateau. Further S the mountain ranges become more isolated
and the deserts between them more extensive. The ‘last refuge’ for a conifer in that direction is
in the Hajar Mountains of Oman. Between there and the high escarpment of the Asir Range in
Saudi Arabia above the E shore of the Red Sea stretches the Arabian Desert. There is consider-
able overlap with other regions such as the Mediterranean in the distribution of conifers and
only species that are endemic to this region or have their major distribution here are treated in
this section.

Pinaceae

Map MAJ-5. Picea orientalis

[n=40, m=39, h=10, 1862–2008]

Picea orientalis occurs around

the E part of the Black Sea
in the Pontus Mountains of
Turkey, the Transcaucasus of
Georgia and in the main Cau-
casus Range from W Georgia
to Dagestan (Russian Federa-
tion). The westernmost popu-
lation is in the Canik Dağları
at 37° 06’ E. It is a forest forming tree occurring in the mountains at altitudes between 700 m and
2100 m. It forms extensive pure stands, especially at higher elevations and at the limit of trees,
or is mixed with Abies nordmanniana and to a lesser extent Pinus sylvestris. At lower elevations
it occurs scattered in broad-leaved forests, with Fagus orientalis, Quercus spp., Acer spp., Ilex
colchica and Taxus baccata.
 mainland asia and japan 177

Map MAJ-6. Abies cilicica [n=32,

m=30, h=9, 1853–2011]

Abies cilicica occurs in the moun-

tains of S Turkey and in Lebanon
and Syria. Its distribution is similar
to that of Cedrus libani var. libani
(map EM-28 on p. 157) but it is rare
outside Turkey so it is treated here
and not in the chapter Europe and
the Mediterranean. In Turkey it is
common in the Taurus Mountains
(Toros Daglari) and in the W part
(Isaurian Taurus) a distinct sub-
species A. cilicica ssp. isaurica is
recognized. The distinction is not
always recognized and we have
not separated them on the map;
there may also be some sympa-
try of the two taxa. Abies cilicica
often occurs with Cedrus libani at
altitudes between 1000 m and 2100 m or it forms pure stands especially at the higher end of its
altitude range. In Syria and Lebanon the original forests have mostly disappeared long ago and
this species remains in isolated stands in forest remnants. The species can be listed as threatened
with extinction in these countries, but a global assessment also considers the still extensive for-
ests in Turkey that contain this species, so it is not listed in a threatened category on the IUCN
Red List.

Map MAJ-7. Abies nordmanniana ssp.

nordmanniana (black) [n=34, m=33,
h=7, 1836-2009]; A. nordmanniana ssp.
equi-trojani (red) [n=24, m=23, h=6,
1883–2005]

Abies nordmanniana is distrib-

uted in the mountains surround-
ing the Asian side of the Black
Sea; it is the only species of fir
in this region. Traditionally, three subspecies are recognized, but distinctions between two of
them are so minor that they are not separated here (Farjon, 1990, 2010a). Abies nordmanniana
ssp. equi-trojani occurs in the Pontus Mountains of N Turkey and westward as far as Kaz Dağ
(Mt. Ida) the mountain in Homer’s Iliad from where the gods ­sometimes watched the battles for
Troy just a few km away, hence the epithet (equi-trojani = of the Trojan Horse). In these W parts
of Turkey the fir forests are isolated on a few of the highest mountains; further E where subspecies
equi-trojani and ssp. nordmanniana may occur in the same area such forests are more frequent,
only to become uncommon again in the Anatolian Mountains (Anadolu Daglari). More extensive
forests of Abies nordmanniana and Picea orientalis occur in Artvin (NE Turkey) and in parts of
the Transcaucasus in Georgia. Isolated from these by a plain that widens out towards the Black
Sea are the largest fir-spruce forests in the Greater Caucasus on both sides of the border between
Georgia and the Russian Federation. Abies nordmanniana occurs at altitudes between 900 m and
2100 m. On the mountains Kaz Dağ and Ulu Dağ in NW Turkey the subsp. equi-trojani extends
to the tree line at 1900–1950 m, where it forms small windswept stands among extensive ground
covering ‘mats’ of Juniperus communis var. saxatilis.
178 mainland asia and japan

Abies nordmanniana ssp. equi-trojani with Juniperus communis var. saxatilis on the mountain Ulu Dağ
(Mt. Olympus) in W Turkey. Photo credit Aljos Farjon.

Cupressaceae

Map MAJ-8. Juniperus excelsa ssp. excelsa (black) [n=96, m=79, h=16, 1700–2009]; J. excelsa ssp. polycarpos
(red) [n=159, m=130, h=15, 1830–2006]

Juniperus excelsa is one of the most widespread species of juniper in Eurasia, extending from
Macedonia, W Bulgaria and Greece in Europe to Kirgyzstan and Tajikistan in Central Asia and
N Pakistan and NW India (a single locality is known from a herbarium specimen at K in Shimla in
Himachal Pradesh; J. semiglobosa has been confused with J. excelsa ssp. polycarpos in literature on
Indian conifers). The northernmost localities are in the Crimea of Ukraine and the southernmost
population occurs in the Hajar Mountains of Oman. Two subspecies are here recognized which
roughly separate in a W and E range of the species as a whole. Juniperus excelsa ssp. excelsa is
the W subspecies, with all European, Lebanese and Iraqi and most Turkish populations belonging
to it. To the E of these, where the climate becomes more continental and arid and the junipers
form high altitude forests, J. excelsa ssp. polycarpos occurs. This taxon has been classified as a
species (or more than one species) in Central Asia but it is best understood as a subspecies of a
widespread and variable species (Farjon, 1993; 2005) even though in such a vast range genetic and
morphological differences may be observed in various mountain systems. The altitude range of
the species is great, from 100 m to 3950 m, with the lowest range in the Mediterranean W and the
highest altitudes reached in Iran (Kerman Province) and in the Karakoram-Himalaya. In Central
Asia, it is restricted to the more western parts of the numerous mountain ranges, where it forms,
with some other species, ( J. semiglobosa, J. pseudosabina, J. sabina) extensive juniper forest or
woodland (‘Archa forest’) on all but the N-facing slopes.
 mainland asia and japan 179

Central Asia and Siberia

Geography

Central Asia (‘Middle Asia’ in Russian literature) is a region in the centre of the Asian continent
including five now independent republics of the former Soviet Union: Kazakhstan, Kirgyzstan,
Tajikistan, Turkmenistan and Uzbekistan. Broader definitions of the term can include several
neighbouring countries, e.g. Afghanistan and Xinjiang Province of China. For this Atlas we use the
term Central Asia as defined above plus Afghanistan. Also included in this section of the chap-
ter is Siberia including European Russia but excluding the far eastern parts i.e. Kamchatka and
Primoriye, and Mongolia. Central Asia, with the exception of the vast steppes of Uzbekistan and
Turkmenistan which extend from the Caspian Sea to the Altai Mountains, is a very mountainous
region, with the Tian Shan system trending E-W and to the S of this the ‘knot’ of mountains con-
necting it with the Himalayan system. Several peaks here exceed 7000 m, creating large glaciers. N
Russia and Siberia are predominantly lowland expanses, with the N-S trending Urals traditionally
separating European Russia from Siberia. Large rivers in Siberia drain northward into the Arctic
Ocean. This whole region is dominated by taiga forest up to the tree line at or N of the Arctic
Circle. The Altai Mountains are situated in the region where China, Mongolia and Russia join bor-
ders, E of these are the Mongolian highland steppes merging with the Gobi Desert. The climate
ranges from Arctic to extreme continental, including desert conditions. Mountains receive higher
precipitation levels, although much of it as snow. Conifers are ubiquitous in the taiga forest and
common in most of the mountain ranges, but taxonomic diversity is limited. It must also be noted
that the distribution of conifer species in the taiga forest of Russia/Siberia is very incomplete on
the maps presented here, as the dots represent herbarium collections only. The main herbarium
consulted in Russia is LE where due to time constraints and the necessity of assisted translation
the collections could only be sampled partly.

Pinaceae

Map MAJ-9. Pinus sibirica [n=47,

m=46, h=11, 1886–2007]

Pinus sibirica occurs in China in

Heilongjiang (Tuqiang), Nei Mon-
gol and Xinjiang, in Kazakhstan,
in Mongolia and in Siberia; an iso-
lated population is reported from
the Kola Peninsula in NW Rus-
sia. In Kazakhstan, Xinjiang and
Mongolia it is only present along the N borders, mainly in the Altai Mountains. No herbarium
collections could be found to substantiate the records for Heilongjiang and Nei Mongol (Inner
Mongolia) in China given in Flora of China (vol. 4, 1999) and for the Kola Peninsula in Russia.
The latter occurrence, if correct, is significant in connection with the present distribution of the
closest relative of this species, P. cembra (map EM-13 on p. 148) which reaches its furthest point
E in the Carpathian Mountains of W Ukraine. Both species have a mutualistic relationship with
the same bird, i.e. the Eurasian nutcracker (Nucifraga caryocatactes) which exclusively disperses
the seeds (Farjon, 2008). This bird occurs in the taiga forest of Russia and Siberia, in Belarus, the
Baltic States and Poland, in southern parts of Norway and Sweden and in the Carpathians, Alps
and Balkan Mountains. In extreme winters some birds from Siberia migrate far into NW Europe.
In colder phases during the Pleistocene this bird could have spread the two pine species beyond
their present and more restricted ranges. Or we could postulate one ancestral species separating
into two that now occur far apart, but still being served by the same more widespread bird.
180 mainland asia and japan

Map MAJ-10. Picea obovata (incl. P. × fennica) [n=58, m=54, h=13, 1827–2003]

Picea obovata is the Russian/Siberian sister species of P. abies in Europe (map EM-17 on p. 151).
Where the two species of spruce meet in Karelia, N Finland, the Kola Peninsula and perhaps the
Komi Republic hybrids known as Picea ×fennica occur; such herbarium records are here included
in the map of P. obovata. This species is the ubiquitous spruce tree of the Siberian taiga, forming
monospecific conifer forests in large parts of its wide range that extends from Karelia and the Komi
Republic to E Siberia. The map, apart from being very incomplete, shows collection bias favouring
the Altai Mountains, where this species crosses borders with China, Kazakhstan and Mongolia.

Map MAJ-11. Picea schrenkiana ssp. schren-

kiana (black) [n=41, m=36, h=8, 1840–1980];
P. schrenkiana ssp. tianschanica (red) [n=22,
m=21, h=5, 1887–1987]

Picea schrenkiana is the spruce tree of

the Tian Shan. The Tian Shan (Celestial
Mountains) is a large system of moun-
tain ranges that includes several ranges
in Kirgyzstan situated to the N of the
main range which extends eastwards
into Xinjiang, China (including Bogda Shan). In Kirgyzstan, P. schrenkiana occurs most abundantly
in the ranges to the N of the main chain, from the Chatkal Range in the W to the mountain ranges
surrounding Lake Issyk Kul. From there it occurs further E into China along the main range of
the Tian Shan as far as Bogda Shan to the E of Urumchi. Its northernmost population is in the
Dzhungarskiy Alatau in Kazakhstan. Two subspecies are recognized, P. schrenkiana ssp. schrenki-
ana and P. schrenkiana ssp. tianschanica; their ranges meet in Kirgyzstan but the latter subspecies
extends further S into the ranges surrounding the upper Naryn River and into the main range of
the Tian Shan in the most E part of Xinjiang. Some distance S of this range are the southernmost
occurrences of P. schrenkiana near Kon-
gur Shan, a mountain 7719 m high. This
species forms pure forests of columnar
crowned trees up to 50–60 m tall on the
slopes and crags at altitudes between
1300 m and 3600 m.

Picea schrenkiana ssp. schrenkiana in Ter-

skey Ala-too, mountains S of Lake Yssyk Kul,
Kirgyzstan. The spruce trees form pure (sin-
gle species) forest from artificial meadows
created for horse grazing in the valley up to
the tree line. Photo credit Aljos Farjon.
 mainland asia and japan 181

Map MAJ-12. Larix sibirica [n=83,

m=78, h=10, 1827–2008]

Larix sibirica is a major com-

ponent of the coniferous taiga
forest and is widespread in
N Russia and W and central
Siberia, the N part of Xinjiang
in China and parts of Mon-
golia. An isolated population
occurs near Lake Onezhskoye in Karelia (not shown on the map) which is the westernmost occur-
rence at ca. 35° E. From there it follows the coast of the White Sea but further E it recedes inland
and meets the mouth of the River Ob on the E side of the Urals. In Siberia it extends as far E as Lake
Baikal. Many herbarium collections were made in the Altai Mountains but few in N Russia and in
much of Siberia. In the E of its range L. sibirica meets L. gmelinii and a zone of hybridization pro-
ducing intermediate forms (L. ×czekanowskii) is said to occur from Lake Baikal and the Lena River
N to the Arctic Ocean. Extensive collecting by Chinese botanists in the period 1950–1970 has shown
the occurrence of L. sibirica in the Altai on the Chinese side of the border and in several locations
in mountains to the S, with the location in Beida Shan at 43° 10’ the most southern point known for
this species. While in European Russia L. sibirica does not form the Arctic tree line, it does so from
the Urals eastwards, with small, stunted trees surviving the long and harsh winters around the Arctic
Circle. In the southernmost reaches of its range (Xinjiang, Mongolia) it is the dominant tree of the
‘forest steppes’ i.e. isolated and often open stands of trees in Artemisia steppe.

Map MAJ-13. Abies sibirica ssp. sibirica (black) [n=49, m=48, h=8, 1826–1998]; A. sibirica ssp. semenovii (red)
[n=18, m=15, h=6, 1897–1995]

The two subspecies of Abies sibirica are geographically separated, with a wide distribution of
A. sibirica ssp. sibirica and a limited distribution of ssp. semenovii. Abies sibirica spp. sibirica is a
component of the Siberian taiga but is also present in the western parts of the Altai Mountains.
This subspecies extends from the Urals in the W to Yakutiya in the E. It remains well south of the
arctic tree limit in Siberia, in fact it is more common in W Siberia and the Altai Mountains, which
have a less severe climate. Abies sibirica ssp. semenovii occurs in the Chatkal Range (Uzbekistan
and Kirgyzstan), in the Talass Ala-too, a parallel range to the N in Kirgyzstan, and in the trans-
verse range Fergana-Too Tizmegi, also in Kirgyzstan. It occurs mainly on N-facing slopes or in steep
ravines at altitudes between 1300 m and 2850 m. Abies sibirica ssp. semenovii is sympatric with
Picea schrenkiana ssp. tianschanica (map MAJ-11 on p. 180) in the Chatkal Range and mountains
surrounding the upper Naryn River to the S, all in W Kirgyzstan. Further E the conifer forests are
182 mainland asia and japan

of Picea ­schrenkiana only. Because P. schrenkiana, while sympatric with A. sibirica ssp. semenovii in
W Kirgyzstan, extends vertically to higher altitude, climate factors probably are the main causes for
this change in the composition of the dominant trees in the conifer forests of Central Asia.

Cupressaceae

The only members of this family in Central Asia and Siberia are species in the genus Juniperus.
Central Asia, especially in its W parts, has extensive juniper forests (‘Archa’ forests) with several
species present. Most of these have wide distributions to Western Asia, the Himalayas, or even
across N Eurasia ( J. sabina map EM-33 on p. 162) or circumpolar ( J. communis maps EM-31–32 on
pp. 160–161) and are given maps elsewhere in this Atlas. Therefore only one species is treated here.

Map MAJ-14. Juniperus pseudosabina

[n=131, m=108, h=17, 1840–2008]

Juniperus pseudosabina is widely

distributed in Central Asia and
eastwards to Xinjiang in China and
to Mongolia. It occurs in Afghani-
stan (Takhar), China (Xinjiang),
Kazakhstan (southern mountains),
Kirgyzstan, Mongolia, Pakistan
(Baltistan, Hindu Kush, Karakoram Range), ­Tajikistan and Uzbekistan (Turkestan Range). In E Kir-
gyzstan it is common in the upper zone of Picea schrenkiana forests; in the W part of its range from
W Kirgyzstan to N Pakistan it forms alpine juniper woodland (‘archa’ forest) with J. semiglobosa
(map MAJ-31 on p. 193) and J. excelsa ssp. polycarpos (map MAJ-8 on p. 178). In Xinjiang and Mongo-
lia it occurs in forested locations with Larix sibirica (map MAJ-12 on p. 181) in ‘forest steppes’ where
it is a decumbent shrub instead of an upright shrub or a small tree in its most western locations.
The altitude range is 1950–4100 m, with the highest occurrences in N Pakistan.

Juniperus pseudosabina in the Alai Range S of Osh, Kirgyzstan. Photo credit Aljos Farjon.
 mainland asia and japan 183

The Himalayas

Geography

The Himalayas here include the Karakoram, Hindu Kush and Himalaya Ranges, forming a con-
tinuous system of folding and uplift caused by the collision of the Indo-Australian and Eurasian
tectonic plates. The Wakhan Corridor in NE Afghanistan separates this chain from the high pla-
teaus of Tajikistan; at the E end the Himalaya Range is bounded by the great bend in the Yarlung
Zangbo-Brahmaputra River; it has a total length of ca. 3500 km. It is the highest mountain range in
the world, with 14 peaks over 8000 m and more than 100 summits in excess of 7200 m. The climate
varies considerably with altitude but also between the S-side and the N-side of the main ranges as
well as generally from E to W. There is a summer monsoon from June to September, with highest
precipitation in the E parts of the range (Arunachal Pradesh, Bhutan, Sikkim, E Nepal) gradually
becoming drier westwards with semi-arid conditions in the inner valleys of the Hindu Kush and
Karakoram. Extensive permanent snow due to altitude and thousands of glaciers provide melt water
continuously even in these drier parts of the range. Almost all conifer species in the Himalayas
belong to families and genera of northern (Laurasian) origin. They belong to the families (and gen-
era) Cephalotaxaceae (Cephalotaxus), Cupressaceae (Cupressus, Juniperus), Pinaceae (Abies, Cedrus,
Larix, Picea, Pinus, Tsuga) and Taxaceae (Amentotaxus, Taxus). The exception is the occurrence in
the E Himalaya of the very widespread species Podocarpus neriifolius, which undoubtedly arrived
here from SE Asia through dispersal by birds. In total, 27 species of conifer occur in the Himalayas.
Cephalotaxus mannii has been found near Satuk, East Kameng District, Arunachal Pradesh, India,
but its wider distribution is in Indochina so this species is treated in that section of this chapter.

Pinaceae

The family Pinaceae is the most diverse in the Himalayas with 14 species present, i.e. four species
in the genus Abies, one in Cedrus, two in Larix, two in Picea, four in Pinus and one in Tsuga. They
are the major constituents of most of the conifer forests in these mountains and occur either pure
(one species in the forest) or in various mixtures, in the latter case with just a few species present
in most below 2500 m, Cedrus below 3300 m, Tsuga below 3300 m, Picea below 3700 m and Abies
and Larix reaching the highest altitudes up to 4100 m. These are the maxima and several species
have substantial ranges of altitude at which they can occur, which also vary with E-W locations
and N-S exposure, so that mixed conifer forests are present in many locations.

Cedrus

A map of the genus is presented in the chapter Europe and the Mediterranean (map EM-26 on
p. 157); one of the species occurs in the Himalayas.

Map MAJ-15. Cedrus deodara [n=46, m=44, h=7,

1856–2008]

Cedrus deodara is distributed in thee western

part of the Himalayas and in mountains S of
the Hindu Kush. In Afghanistan it occurs from
Urgun in N Paktika through Paktia and Low-
gar to the Hindu Kush in Nurestan, continuing
in that range in the Northwest Province State
of Pakistan and turning SE through Jammu &
Kashmir to the Himalaya. In the NW of their
distribution range the cedar forests stay well
away from the Karakoram and Ladakh Ranges,
184 mainland asia and japan

but further E they enter the main Himalaya, even crossing the border into Xizang [Tibet] in
one location (not shown for lack of herbarium record). Cedrus deodara grows in a belt at eleva-
tions between 1700 and 3000 m a.s.l. in the W part of its range and between 1300 and 3300 m in
the E part, where the climate is less dry. The easternmost distribution is limited by increasing
monsoonal influence, but extremes of dryness and cold in the valleys of the Karakoram Range
and Pamir Knot have halted its expansion to the N. At higher elevations it forms a coniferous
forest belt with, among other species, Abies pindrow, A. spectabilis in Nepal, Pinus wallichiana,
Picea smithiana, and Cupressus torulosa, but forms often also pure stands. As is the case with the
Mediterranean cedars, the Himalayan cedar forests have great cultural significance for the people
who live in these mountains.

Cedrus deodara in the Shimla region, Himachal Pradesh,

India. Photo credit and © Dendrological Atlas Team (Zsolt
Debreczy & István Rácz).

Pinus

Five species of pine occur in the Himalayas, Pinus armandii, P. bhutanica, P. gerardiana,
P. roxburghii and P. wallichiana; P. gerardiana and P. roxburghii belong to subgenus Pinus and the
other species to subgenus Strobus. Pinus armandii has its widest distribution in China and Taiwan
and is treated there in this chapter.

Map MAJ-16. Pinus gerardiana [n=28, m=25, h=12,

1847–1993]

Pinus gerardiana is distributed in the moun-

tains of E Afghanistan, in Pakistan (mountains
in N Baluchistan, the Hindu Kush, Karakoram
and Himalaya ranges) in India (Himalaya in
Jammu-Kashmir and Himachal Pradesh) and
in China in the valley of the Xangquan River
(Langqen Zangbo) in extreme SE Xizang [Tibet].
It occurs in relatively dry valleys where much of
the annual precipitation falls as snow in win-
ter. It occurs at altitudes between 2000 m and
3350 m which means it is mostly confined to the lower parts of valleys in the mountains, where
it tends to prefer dry, sunny slopes. The edible seeds are harvested by local people and in several
 mainland asia and japan 185

areas trees are also planted for seeds, which could mean that distinction between natural occur-
rence and introduction has become uncertain, e.g. in the more southern locations in Afghanistan
and in Baluchistan. The occurrence in the Himalaya ranges of NW India is under-represented in
the map as a result of few available herbarium collections.

Pinus gerardiana above the Satluj

River in the W Himalaya, Himachal
Pradesh, India, at ca. 1900 m. Photo
credit and © Dendrological Atlas
Team (Zsolt Debreczy & István
Rácz).

Map MAJ-17. Pinus roxburghii [n=36,

m=34, h=11, 1800–2000]

Pinus roxburghii is distributed

along the greater length of the
Himalaya Range from Chitral in
N Pakistan to the West Kameng
District of Arunachal Pradesh, NE
India. It is widespread and com-
mon in the north-south oriented
outer valleys of the Himalaya and
its foothills and often forms pure stands especially on dry, fire-prone slopes. The available her-
barium collections are insufficiently abundant to show this on the map and apparent gaps in the
distribution are due to this lack of data. Its altitudinal range is from 400 to 2300 m a.s.l., with the
highest growing, scattered individuals at 2500 m. Pinus roxburghii occurs in a lower coniferous or
mixed forest belt, below the higher zone dominated by Abies and Larix. Its most common associ-
ated conifer species are Cedrus deodara and Pinus wallichiana, but often it forms light pine woods
in which it is the dominant tree. This species is of considerable biogeographical interest because
it appears to be closely related to some Mediterranean pines and especially to P. canariensis (map
EM-16 on p. 150). Evidence from the fossil record (Klaus, 1988) suggests that these pines were
more widespread in the Cenozoic and that their ranges may once have been connected.

Map MAJ-18. Pinus wallichiana [n=74,

m=73, h=13, 1824–2009]

Pinus wallichiana is the most

common and widespread species
of pine in the Himalayas, occur-
ring along the entire length from
the Hindu Kush in Afghanistan to
beyond the Himalaya proper into
SE Xizang [Tibet], far NW Yunnan
186 mainland asia and japan

and the Nu (Salween)‒Nmai divide in E Myanmar (Burma). As with several other conifers in
the Himalayas, there is a paucity of herbarium collections in some regions, notably Arunachal
Pradesh in NE India where access has long been denied to ‘foreigners’ because of border tensions
between India (which controls the region) and China (which claims it belongs to that nation).
From the West Kameng District in Arunachal Pradesh var. parva has been described; it may also
occur in adjacent Xizang [Tibet] but too few specimens are available to map it separately and it
appears to occur within the eastern part of the distribution of var. wallichiana. Pinus wallichiana
grows in the valleys and foothills, to a maximum altitude of 2700 m, but in Bhutan it reaches
3400 m a.s.l. Sometimes it forms pure stands, in other places it appears as an important forest
component mixed with angiosperm trees, e.g. species of the genera Quercus, Acer and Ilex. In the
W Himalayas, where conditions are drier, P. wallichiana forms mixed forests with Cedrus deodara.
Other conifers with which it may be associated are Pinus roxburghii, Abies spectabilis, or A. densa
and Tsuga dumosa in the wetter E part of its range.

Pinus wallichiana in the Langtang

region of Nepal, Central Himalaya, at ca.
2900 m on a ridge in the middle distance
(the conifer in the foreground is Tsuga
dumosa). Photo credit and © Dendrolog-
ical Atlas Team (Zsolt Debreczy & István
Rácz).

Map MAJ-19. Pinus bhutanica [n=17,

m=17, h=7, 1938–2006]

Pinus bhutanica was first des­

cribed and named as distinct
from P. wallichiana from collec-
tions made in Bhutan in June
1979. Very few earlier herbarium
collections have subsequently
been identified as this species,
among which is one by K. C. Sahni
in 1977 from the Tenga Valley in Arunachal Pradesh and another by K. D. Rushforth in the same
year at the Dongjug—Yigong Zangbu confluence in Bomi Xiang, Xizang [Tibet]. Another location
reported in Arunachal Pradesh (Lohit Valley) is not mapped for lack of herbarium specimens. In
the 1980s some further collections were made of this pine in Bhutan and, more recently, in Gong-
shan Xian in the far NW of Yunnan by Sino-American biodiversity surveys (specimens at CAS),
where it was found in several localities between the Dulong Jiang and Nu Jiang (Salween). This
history of collection efforts and the gap between localities in Bhutan and NW Yunnan strongly
indicate its likely presence in Arunachal Pradesh where, as remarked above, few botanists have
travelled. The species occurs in a (warm) temperate zone, from ca. 1700 m to 3050 m in Bhutan,
at 2100 m in SE Xizang [Tibet], and at 1800–2200 m a.s.l. in NW Yunnan, with reports down to
1000 m in Arunachal Pradesh. Associated species can include Pinus wallichiana, Pinus roxburghii
(dry inner valleys) and various angiosperm trees.
 mainland asia and japan 187

Pinus bhutanica in the Tangmai region

of Xizang [Tibet], China at 2600–3000
m mixed With Cupressus torulosa
(darker trees) and scattered P. densata
and Tsuga dumosa. Photo credit and
© Dendrological Atlas Team (Zsolt
Debreczy & István Rácz).

Picea

Map MAJ-20. Picea smithi-

ana (black) [n=24, m=21,
h=13, 1856–2009]; Picea spinu-
losa (red) [n=21, m=20, h=5,
1849–1986]

The genus Picea is rep-

resented in the Himala-
yas by two species [Picea
brachytyla is reported
from Arunachal Pradesh
by Sahni (1990: 74) but no
collection could be found at K] and both are limited to these mountain ranges, but separated in
their distribution. Picea smithiana is the westernmost species and occurs from Konar in Afghani-
stan to Central Nepal and the Gyirong Valley in Xizang [Tibet]. Picea spinulosa occurs in Sikkim,
Bhutan, Arunachal Pradesh and adjacent parts of Xizang [Tibet]. The available herbarium col-
lections of both species are rather limited in number, therefore the distribution shown is incom-
plete and apparent gaps in the Himalaya of Himachal Pradesh and Uttar Pradesh of India for P.
smithiana and in Arunachal Pradesh for P. spinulosa are most likely artefacts of insufficient col-
lecting in these areas. The gap between the two species in E Nepal is a real one, given the relative
abundance of collections from either side and the fact that E Bhutan has been well visited by
collecting botanists for a long time. Spruces do not occur there, it appears. The altitude range of
both species is similar, ca. 2400–3700 m, and both occur in mixed coniferous forests with mostly
the same associated co-dominant species (but Cedrus deodara and Abies pindrow only with P.
smithiana and Abies densa only with P. spinulosa). The gradually drier climate from E to W only
becomes significant W of Nepal, so this neither explains the gap in E Nepal nor the absence of P.
spinulosa in that country.

Larix

Two species of Larix are native in the Himalayas and both are variable species which have led
some authors to recognize more species. These taxa are here recognized as varieties. The genus
Larix has a northern boreal distribution and the Sino-Himalayan mountain system is the only
region where some of its species reach more southern latitudes (map GTC-20 on p. 26). In the
Himalayas, the larches are confined to the E half of the mountain range where monsoon influ-
ence is strongest.
188 mainland asia and japan

Map MAJ-21. Larix griffithii var. grif-

fithii (black) [n=52, m=51, h=7, 1838–
2001]; L. griffithii var. speciosa (red)
[n=15, m=15, h=6, 1924–2000]

Larix griffithii var. griffithii occurs

in the Himalaya from N-Central
Nepal to Arunachal Pradesh (in
the latter region of India few her-
barium records are available) and
also on the Tibetan side. The spe-
cies is also present away from the
Himalaya in SE Xizang [Tibet], in NW Yunnan and in N Myanmar (Burma) but we have only
George Forrest’s collections from the border with Yunnan (W side of divide) for that country;
these belong to var. speciosa. In SE Xizang var. griffithii is joined by var. speciosa and the latter
prevails in Yunnan, so there is geographic separation to some extent. Larix griffithii var. griffithii
is apparently the Himalayan taxon and occurs much further to the W than var. speciosa. Unfor-
tunately the paucity of herbarium collections from Arunachal Pradesh and Myanmar (Burma) do
not allow us to have a wider view of where the two taxa occur in the same area and where they
separate. According to Sahni (1990) L. griffithii occurs continuously from “E Nepal through Darjeel-
ing, Sikkim, Bhutan, Arunachal Pradesh, NE Upper Burma, and [in the] Chumbi Valley in Tibet”
and L. speciosa is not mentioned for the Himalaya. The Chumbi Valley is situated between Sikkim
and Bhutan, but L. griffithii var. griffithii is known from many other localities on the Tibetan side
of the Himalaya. Larix griffithii occurs in the cloud belt of the Himalayas, at elevations between
(1800–)2400–4000(–4100) m, commonly on rocky moraines. It occurs in pure forests up to the tree
line, at lower elevations it is often mixed with Abies spectabilis, A. densa, Pinus wallichiana, Picea
spinulosa and Tsuga dumosa.

Map MAJ-22. Larix potaninii var. himalaica

[n=7, m=7, h=4, 1975–2001]

Larix potaninii is a variable species almost

exclusively occurring in China (see map
MAJ-63 on p. 215) but one variety, L. pota-
ninii var. himalaica, occurs in a small area
of the Himalaya, on the Nepalese and on
the Chinese (Tibetan) side of the bor-
der and we present a map of it here. It
is known from a limited number of her-
barium collections and some of the stated
toponyms are difficult to locate, so some
dots on the Chinese side of the border are
approximate localities. They are all in the
Gyirong River Valley and some tributary
streams, a trans-Himalayan river which
leads down across the border to become
the Trishuli River in Nepal. The Nepalese localities of L. potaninii var. himalaica are in the valley
of the Langtang Khola, an eastern tributary of the Trishuli. The herbarium collections were made
at altitudes between 3100 m and 3800 m. This variety is listed as Near Threatened (NT) on the
IUCN Red List; only the Nepalese population is in a national park.
 mainland asia and japan 189

Tsuga

Map MAJ-23. Tsuga dumosa [n=116,

m=110, h=14, 1821–2005]

Tsuga dumosa is distributed in

the Himalaya from Uttarkhand
(India) to the Yarlung Zangbo-
Brahmaputra Bend and beyond
to the E of this range in SE Xizang
[Tibet], N Myanmar (Burma),
NW Yunnan and SW Sichuan; a disjunct locality where this species occurs is Mt. Fan Si Pan
in Lao Cai Province, N Viet Nam. Herbarium specimens are nearly absent from Arunachal
Pradesh, but T. dumosa is as widespread there as elsewhere in the E Himalaya. More peculiar
is the absence on the map of records from W Nepal. Presumably it becomes less common there
as the wet monsoon influence diminishes, but it must be present, yet we have found no her-
barium data. When all species of conifer that occur across this region are mapped (not shown)
from the Conifer Database, both W Nepal and adjacent Uttarkhand have markedly fewer dots
on the map than the Himalaya E and W of it. This indicates that the region is under collected
for conifers. Tsuga dumosa occurs in a high belt of conifer forest at altitudes between 2600
m and 3200 m in the Himalaya (1700–3500 m in China) and often accompanies Abies, Picea,
Larix and Pinus as a shade tolerant tree below the canopy, or in pure stands on very wet,
N-exposed slopes.

Abies

Of the four species of Abies present in the Himalayas, A. delavayi is known from a few specimens
collected in Arunachal Pradesh but its main range is in China, so it is treated there. This leaves
A. densa, A. pindrow and A. spectabilis as the three truly Himalayan firs.

Map MAJ-24. Abies densa [n=29,

m=26, h=5, 1881–1995]

Abies densa occurs in the eastern-

most part of the Himalaya from
the Barun Khola near Mt. Makalu
in E Nepal to Motuo Xian in SE
Xizang [Tibet]. Herbarium col-
lections from Arunachal Pradesh
are very rare because the region
has long been kept closed to visi-
tors. This species undoubtedly
occurs there, so the map remains
incomplete. This species occurs in
a wide ­altitudinal range between
2150 m and 4100 m from mixed deciduous coniferous forest at lower elevations to stands with
Betula utilis at the tree line. In the ‘cloud belt’ around 3000–3500 m it often is the dominant tree
in very moist conifer forest on steep rocky slopes and on exposed ridges.
190 mainland asia and japan

Map MAJ-25. Abies spectabi-

lis (black) [n=27, m=26, h=7,
1832–2009]; A. pindrow (red)
[n=32, m=29, h=5, 1836–2008]

Abies spectabilis and A.

pindrow have overlapping
general distributions across
the Himalayas from the
Hindu Kush in Afghani-
stan to Nepal. We map
them together here for this
reason to show the shift
in occurrence from E to
W, but in the W the herbarium records are incomplete for A. spectabilis which occurs as far as
Afghanistan accoding to the literature (Liu, 1971; Farjon, 1990). Abies pindrow reaches W Nepal
but is further E replaced by A. spectabilis which in turn is replaced by Abies densa in Sikkim,
Bhutan and Arunachal Pradesh (map MAJ-24 on p. 189). Abies spectabilis is evidently uncommon
in the W beyond Jammu & Kashmir as the herbaria consulted had very few specimens identified
as this species and none had adequate locality data to place them on the map. Apart from this
geographical shift between the two species, there is also an altitudinal gradient, with A. pindrow
commonly occurring at a range of 2000–3300 m and A. spectabilis somewhat higher between
2500–3800(–4000) m. Where the two species occur in the same area A. spectabilis commonly
occurs above A. pindrow. Both are forest forming species, occurring either in pure stand or mixed
with other conifers.

Abies pindrow in the Simla

region of Himachal Pradesh,
India, mixed with sporadic
Cedrus deodara, Picea smithi-
ana, Pinus wallichiana and
Taxus contorta. Photo credit
and © Dendrological Atlas
Team (Zsolt Debreczy & Ist-
ván Rácz).

Cupressaceae

Two species of Cupressus and seven species of Juniperus occur in the Himalayas. The two species
in Cupressus have their main distribution here; of Juniperus only J. indica is almost exclusively
Himalayan and the other species extend either to the mountains of Central Asia or into those of
China. Of these, J. recurva and J. squamata have a mainly Sino-Himalayan distribution, with the
latter extending across China as far E as Taiwan. Juniperus semiglobosa occurs in the W parts of
the Himalayas and extends beyond the Wakhan Corridor into the mountains of Central Asia. The
other three species have ranges mostly outside the Himalayas and are treated elsewhere.
 mainland asia and japan 191

Cupressus

Map MAJ-26. Cupressus cashmeriana

[n=48, m=30, h=10, 1838–2001]

Cupressus cashmeriana was first

known from plants and seeds col-
lected by William Griffith in Bhutan
in 1838 and brought to Europe, where
in Paris E. Carrière who described and
named it, working with cultivated
plants sourced from England, got its
provenance wrong and reported it to
be native in Kashmir (Farjon, 1994).
This species is quite widely cultivated
in the E Himalaya and native popu-
lations are hidden in remote areas,
mostly in Bhutan but also in Arunachal Pradesh to the E. In Nepal and Sikkim it is only known from
planted trees. It is sometimes difficult to discern from information on herbarium labels whether
the tree from which the specimen was collected grew in natural habitat, while some planted trees
may not have been far removed from an indigenous population. This influences the accuracy of our
map, which is the best estimate of natural distribution we are able to give based on these data. In
January 2001 Sabine Miehe (Univ. of Marburg, Germany) found trees of C. cashmeriana growing as
emergents on a steep slope in undisturbed evergreen oak forest in Bhutan, Kuru District, along the
lower Yangri Chu N of Tergang which were estimated to be 85–95 m tall. If correct, these would
be the tallest trees growing in Asia. The natural habitat, now often disturbed, is evergreen forest,
fagaceous/lauraceous at lower altitudes and coniferous at higher altitudes, and limestone cliffs and
slopes, at altitudes between 1250 m and 2670 m.

Map MAJ-27. Cupressus toru-

losa var. torulosa (black) [n=51,
m=45, h=12, 1800–2006]; C. toru-
losa var. gigantea (red) [n=20,
m=18, h=7, 1934–1997]

Cupressus torulosa has a dis-

junct distribution, with two
main areas: the W Hima-
laya from Himachal Pradesh
(Chamba) to West-Central
Nepal (Mustang District)
and E Xizang [Tibet] in the
lower Yarlung Zangbo Valley
and tributaries from the N, from near Gyangze downstream to Bomi on the Nagong River. Two local-
ities are known from herbarium collections in between these two areas, one from W Bhutan and
one from Arunachal Pradesh, India. The latter region has seldom been visited by botanical collectors
as the region has been inaccessible for a long time for political reasons. The Tibetan populations
are considered by Chinese botanists to belong to C. gigantea, here recognised only as a variety of
C. torulosa (Farjon, 2005, 2010a) and it is obvious from observations during site visits by Sabine Miehe
(pers. comm.) and the collected specimens that the two forms occur in the same area. The near
absence of this species in the Himalaya from West-Central Nepal to E Xizang [Tibet] is likely caused
by the monsoon influence, which diminishes from E to W across the range. Cupressus torulosa is
192 mainland asia and japan

confined to the drier valleys in the inner parts of the Himalaya, where it occupies S-facing slopes
at altitudes between 1560–3670 m and where annual precipitation does not exceed 300 mm. The E
Himalaya is mostly too wet for this species, as are the more southern valleys in the range in W Nepal.
North of the Himalaya, in the Tibetan valleys that drain south through the Himalaya chain or north
and east into theYarlung Zangbo system, conditions are too dry, but eastwards along the Zhangbo
River downstream from Gyangze C. torulosa is again found in isolated stands. Cupressus torulosa
var. gigantea is listed as Vulnerablee (VU) on the IUCN Red List. Habitat degradation is the most
serious threat.

Juniperus

Map MAJ-28. Juniperus squamata [n=156, m=142, h=21, 1800–2005]

Juniperus squamata has a very extensive distribution, from Afghanistan to Taiwan. It is mostly a
high altitude species with a shrubby habit and is most widespread in the Karakoram–Himalaya
chain and in the high mountains of China in S Gansu, Sichuan and Yunnan. A few localities are
also known in the Hindu Kush (Afghanistan) and in Xizang [Tibet]. All these localities are at
high altitudes, commonly above 3000 m and to a maximum of 4850 m. In Taiwan, it is also a high
altitude species, collected between 2900–3990 m according to our herbarium records. In E China
J. squamata occurs sporadically in the mountains, but often at lower altitudes, with some records
in Anhui, Fujian and Shaanxi from 730–1340 m. No habitat notes are available for these localities
and they may have been disturbed areas with secondary vegetation. Its most common habitats
are subalpine scrubland or tree line and alpine meadows and rocky areas close to the perpetual
snow line. In Xizang [Tibet], Sichuan and Yunnan J. squamata often grows together with J. pingii
var. wilsonii (map MAJ-103 on p. 239) and the two taxa are sometimes confused in herbaria as
they are morphologically similar.

Map MAJ-29. Juniperus indica var.

indica (Black) [n=124, m=112, h=15,
1838–2009]; J. indica var. caespitosa
(red) [n=27, m=23, h=4, 1952–2001]

Juniperus indica is distributed

along the Himalayan chain from
Hunza in Pakistan to at least Bhu-
tan and possibly into Arunachal
Pradesh, a region from where
we have no records. It occurs in
Xizang [Tibet] from around Lhazê
 mainland asia and japan 193

on the Yarlung Zangbo River E to the valleys of the Nu (Salween) and Lancang (Mekong), mainly
in valleys between the Himalaya and the Yarlung Zangbo, but with a few populations to the N
of that main river drainage. Its easternmost localities are in NW Yunnan in mountains (divides)
between the three great rivers which come from the Tibetan Plateau and run parallel through
very deep valleys there. Juniperus indica is commonly an erect shrub or a small tree, but a low
and often decumbent form to 1 m tall has been found in several localities, sometimes in associa-
tion with the tree form; this has been described as var. caespitosa. It occurs in Nepal and Bhutan
and possibly elsewhere. Juniperus indica is a high altitude species occurring from 3600 m to 4800
m a.s.l. in coniferous forest below or at the tree line, as well as above the tree line where it often
forms rhododendron-juniper thickets together with J. squamata or as the only species of juniper
present. At the highest altitudes it is confined to S-facing slopes.

Map MAJ-30. Juniperus recurva

var. recurva (black) [n=102, m=91,
h=15, 1803–2005]; J. recurva var.
coxii (red) [n=21, m=19, h=7,
1838–2006]

Juniperus recurva occurs in

the E Himalaya, in SE Xizang
[Tibet] and in NW Yunnan
and adjacent parts of Myan-
mar (Burma). It is most com-
mon in Central and E Nepal and W Bhutan and in NW Yunnan, but a general lack of records
from Arunachal Pradesh due to political factors together with known localities in adjacent
Xizang [Tibet] could indicate a third area where this species is quite common. Juniperus recurva
var. recurva occurs throughout the range of the species, while var. coxii is known from E Nepal
to Yunnan. The westernmost locality of J. recurva supported by a verified herbarium record is
in the Kumaun Himal of Uttar Pradesh, India, just W of Nepal. Juniperus recurva is rare in W
Nepal, where it is replaced by J. indica and J. squamata. Unlike most other species in the genus,
J. recurva and especially the variety J. recurva var. coxii with its pendulous foliage and longer
leaves is adapted to a moist forest environment. The monsoon influence being strongest in the
E of the Himalaya and in NW Yunnan therefore explains its general distribution. This species is
usually a tree, with var. coxii when growing in evergreen rainforest capable of attaining 40 m, but
becomes a shrub near the tree line. Its altitude range is 2000–4500 m. The pendulous form (var.
coxii) is often planted in Yunnan and elsewhere and such occurrences when sampled for herbaria
have been excluded from the map.

Map MAJ-31. Juniperus semiglobosa [n=178,

m=132, h=20, 1700–2004]

Juniperus semiglobosa occurs from

Nepal to Kirgyzstan in the Himalaya,
Karakoram, Hindu Kush, Pamir, Alai
and Kirghiz Ranges. This species is
well adapted to dry conditions, which
explains its general western distribu-
tion, but it is rare in the Pamir Knot, the
central mountains, plateaus and high
valleys in Tajikistan, the driest part of
this vast complex of high Asian moun-
tain ranges. It is also absent in the Tien
194 mainland asia and japan

Juniperus semiglobosa in the Ala-

Archa Gorge, Kirghiz Range, Kir-
gyzstan, here growing on a coarse
gravel and boulder river terrace but
also present on the slopes behind.
Photo credit Aljos Farjon.

Shan Range, which extends from

S Kirgyzstan far eastward into
China. Although this range rises
from deserts, its altitude and
perpetual snow and ice provide
water to sustain a forest belt of
spruces (Picea schrenkiana, map
MAJ-11 on p. 180) and junipers
( Juniperus pseudosabina, map MAJ-14 on p. 182) but J. semiglobosa has not been found there.
In the central and SE parts of its range J. semiglobosa occurs in mixed coniferous forest with
Abies, Cedrus and Pinus and above the forest zone in subalpine scrubland on S-facing slopes or
in juniper stands on glacial moraines. In Kirgyzstan it is uncommon in the spruce forests but
forms extensive juniper forests above the spruce forests, or in the W of the country above mixed
angiosperm forest, together with J. excelsa ssp. polycarpos (map MAJ-8 on p. 178) and sometimes
other species e.g. J. pseudosabina or J. sabina.

Taxaceae

Two species of Taxus occur in the Himalayas, T. contorta and T. wallichiana. Taxus contorta,
although described in the mid-nineteenth century by W. Griffith, was subsumed into T. wallichi-
ana and only recently recognized as a distinct species, ‘T. fuana’, whereby Griffith’s long forgotten
name and description were overlooked. Taxus wallichiana occurs in the Himalaya E of T. contorta
and the transition zone is in E Nepal, but its distribution extends far beyond across SE Asia and
into Malesia. Because it is important to show where the two species meet, both are treated here.
One species of Amentotaxus occurs in the far E of the Himalaya chain.

Taxus

Map MAJ-32. Taxus contorta

[n=57, m=44, h=4, 1840–2008]

Taxus contorta is distributed

from Paktia in Afghanistan
through the Hindu Kush
and Himalaya as far as Cen-
tral Nepal around 85° 20’
E. There is a long distance
between T. contorta and
T. baccata (map EM-42 on
p. 169) of which the farthest
eastern record is in the
Elburz Mountains of N Iran.
The intervening mountains
 mainland asia and japan 195

are too dry, as most of Iran and Afghanistan is desert or semi-desert. Winter low temperatures
are another limiting factor for yews, so there are none in the mountains of Central Asia, which
otherwise might form a pathway of migration. In wetter times during the Pleistocene the two spe-
cies may have had contact, but evidence is lacking. Taxus contorta occurs in ­coniferous or mixed
forests at altitudes between 1800 m and 3200 m (rarely in subalpine thickets of Rhododendron up
to 3400 m) and like all yews is shade tolerant and thrives best under the canopy of other trees
unless growing in a cool maritime climate. Exploitation of bark and leaves to extract alkaloids for
the production of anti-cancer drugs has caused very serious decline of this species in large parts
of its range. It is listed as Endangered (EN) on the IUCN Red List.

Map MAJ-33. Taxus walli-

chiana [n=104, m=94, h=16,
1831–2001]

In the Himalaya, Taxus

wallichiana occurs E of
T. contorta in the wetter
parts of the range; the
transition zone between
the two taxa in Central
Nepal more or less coin-
cides with the climatic
transition. It is known
from E Nepal, Sikkim,
Bhutan and Arunachal
Pradesh within the
Himalayan range. How-
ever, T. wallichiana has a
much wider distribution.
In India it also occurs in
the Khasi Hills of Megha-
laya and in the Naga
Hills of Assam. To the E
it occurs in Myanmar, N Yunnan and S Sichuan, to the SE it has been found in Lam Dong and
Khanh Hoa provinces, S Viet Nam. Further disjunct populations occur in Sumatera and Sulawesi
(Indonesia) and in Luzon and Mindanao (Philippines). In Sumatera and Sulawesi this Northern
Hemisphere genus has crossed the equator; indeed the disjunct occurrences in S Viet Nam and
Malesia may be interpreted as long distance dispersal events brought about by migrating birds.
These are chance events and therefore explain absence of yews in other localities with suitable
habitat, e.g. Mt. Fan Si Pan in N Viet Nam and Mt. Kinabalu in Sabah, Borneo, where T. wallichi-
ana would be expected to still occur if range contraction was the cause of the disjunct distribu-
tion. This species is mostly montane to high montane, at altitudes between 900 m and 3700 m,
and occurs as a sub-canopy tree in evergreen forest, either dominated by conifers, mixed, or by
angiosperms. Heavy exploitation of bark and leaves to extract alkaloids for the production of
anti-cancer drugs has caused very serious decline of this species in mainland Asia. It is listed as
Endangered (EN) on the IUCN Red List.
196 mainland asia and japan

Amentotaxus

Map MAJ-34. Amentotaxus assamica [n=4,

m=4, h=3, 1874–1986]

Amentotaxus assamica is a rare species

of which only a few herbarium collec-
tions have been made in the NE Indian
state of Arunachal Pradesh. This region
is disputed territory, administered by
India but claimed by China; as a result it
was inaccessible for many years but now
slowly opening up to visitors. This spe-
cies, though rare, undoubtedly occurs in
more localities than shown on the map.
Amentotaxus assamica occurs in warm
temperate montane rain forest on N-facing slopes at altitudes between 1600 m and 2000 m. These
forests are described as ‘mossy forest’ dominated by Quercus, Castanopsis, Acer and Rhododen-
dron, with associated taxa such as Magnolia, Michelia, Corylopsis himalayana, Betula alnoides,
Carpinus viminea and Exbucklandia populnea. Amentotaxus assamica is listed as Endangered (EN)
on the IUCN Red List due to its rarity and threats from deforestation.
 mainland asia and japan 197

China and Taiwan

Geography

China is after Russia and Canada the third largest country in the world with a land area of
ca. 9.63 million km² (marginally larger than the USA with 9.37 million km²). Its greatest extent
is E-W across the E Asian landmass over a distance of just over 5000 km. In the E China borders
on coastal seas and straights that open into the Pacific Ocean. Two large islands are situated in
the South China Sea (Hainan) and between this and the East China Sea (Taiwan) and numerous
offshore islands fringe the coast. The most general and striking geographical feature of China is
a division into a western and an eastern (smaller) part, the W continental, high, cold and arid,
the E maritime, low, warm-temperate and moist. With ca. 1.34 billion people (census 2010) China
has the largest population of any country but almost all live in the E part where agriculture
is intense and has existed for millennia. As a result of this intensive use only 5.9% of China’s
forest cover is considered old growth or primary forest. Within this bipartite division Chinese
geographers recognize five macro-regions: Northeast Plain, North Plain, Southern Hills, Xinjiang-
Mongolia and Tibetan Highlands. The topographical diversity of China is great, with almost all
possible landforms present, from the deepest desert basins to the highest mountains, with vast
plains at near sea level to the high Tibetan Plateau, numerous long rivers that traverse much of
the country, some of the largest karst landscapes in the world, extensive deserts and vast areas
under cultivation yielding several crops per year, numerous isolated mountain ranges in the SE
interspersed with broad and fertile valleys, snowy ranges away from the Himalayan chain with
peaks higher than any in the Andes sending glaciers down to well below the tree line, boreal
conifer forests in the far NE and tropical rainforest in Hainan Island. What follows is the briefest
possible description of the five regions of China. Debreczy & Rácz (2011 vol. 1: 225–264) give a
more extensive account of a wider part of Asia that includes mainland China and is relevant to
conifer distribution.
In the NE beyond the Changbai Shan (mountains on the border with North Korea) stretches
the vast Manchurian Plain. From E to W this plain changes from boreal lowland forest to grass
steppe and, beyon the Greater Hinggan Range, to the Gobi Desert. The North Plain forms a rough
triangle with Beijing at its N apex, Shanghai in the SE and Yichang in the SW. It is a low alluvial
plain fed by the Yellow River and Yangtze River (the latter 6300 km long and both entirely in
China) with a very dense population and intensive agriculture; only a few isolated mountain
ranges interrupt it. The Southern Hills region is extremely diverse. In the NW lies the Sichuan or
Red Basin surrounded by mountains on all sides and also densely cultivated and populated. S of
the Yangtze River the topography is complex, with continuously mountainous terrain in the W
(often called western China but really part of the E half of the country) and numerous isolated
ranges, often rising steeply from level plains, in the E. In the far S of the country steep karst for-
mations (‘sugarloaf ’ mountains) interspersed with flat bottomland form mazes over more than
100,000 km² extending across the Vietnamese border and in some places continuing into the
South China Sea as islands and sea stacks. Xinjiang-Mongolia is the largest of the five regions. In
the E lies the Loess Plateau, at 650,000 km² the largest of its kind in the world, covering large parts
of Gansu, Shaanxi and Shangxi provinces and creating dust storms as well as causing the colour
of the Yellow River. High mountains and plateaus fringe the Gobi Desert, which extends N into
Mongolia. NW of the Tibetan Plateau, between the Kunlun Shan—Altun Shan and the Tian Shan
lies the vast Tarim Basin, a cold and arid sink into which numerous rivers disappear and with
a deepest point at –154 m. The Takla Makan Desert, one of the driest deserts in the world and
covered in sand dunes, lies in the W part of this basin. From this basin the land rises to the Pamir
and Karakoram Ranges in the W and the Tian Shan in the N, with peaks over 7000 and 8000 m.
The Tibetan Highlands include the Tibetan Plateau, at 2.5 million km² and an average elevation
of >4,000 m the largest and highest plateau in the world (‘Roof of the World’). To the W and S of
this plateau China is bordered by the Himalayan Ranges (see under that section in this chapter).
198 mainland asia and japan

The E to SE section of the plateau is dissected by several large rivers, three of these converge
in NW Yunnan into the deepest parallel mountain valleys in the world. The Tibetan Highlands
extend far E-SE into Sichuan and Yunnan in a series of mostly N-S oriented ranges, culminating
in Gongga Shan in W Sichuan (7556 m).
With such diversity in geography it is not surprising that climate is also diverse and it ranges
between extremes, from subarctic to tropical, from high alpine to summer hot lowland and from
extreme desert to wet monsoon. The Pacific monsoon influence from late spring through the
summer reaches from the E coast (typhoons) far into the mountainous interiors of Sichuan and
Yunnan and well into SE Xizang [Tibet]. The island of Taiwan has some of the highest levels of
rainfall in the world on its mountainous E coast; the island of Hainan is tropical. In much of low-
land E and S China summers are hot and moist while N of the Yangtze winters are usually cold
and dry. In the W of China high mountain ranges block any oceanic influences and perpetual
snow and ice are almost the only sources of water to the interior. On the high Tibetan Plateau
this has resulted in numerous lakes but lower down in the basins and deserts rivers sink into the
sands or evaporate creating salt flats. The topography of E China and of lands to the S has been
conducive to N↔S migrations of plants and animals with the oscillations of climate during the
Pleistocene. This has meant that much of the Cenozoic Holarctic flora still exists in China, while
many species went extinct elsewhere, particularly in Europe. The flora of China is extraordinary
diverse, with an estimated 31,200 species of vascular plants (Flora of China 1, in press) and new
species continuously being found.

Conifers in China and Taiwan

Families Genera Species Species + infra Countries with Area of occupancy Endemic taxa
conifers in km²
5/8 30/70 142/615 222/794 1/1 55,400/9,633,000 124
Numbers after ‘/’ are global or continent totals.

Given the size of the country, the extraordinary variation in topography and climate and the
overall size of its flora, it is not surprising that China has a very diverse conifer flora as well. In
the above calculations Taiwan is included, which has many endemic taxa of its own and is treated
in a distinct section after mainland China. At five, the number of families present is one less than
for the chapter, as Sciadopityaceae is endemic to Japan. The number of genera, 30 or 42.9% of
the world total, is very high, compared to e.g. the USA with only 17 genera. The number of spe-
cies, though slightly higher, is more similar to the USA. Both countries have similar total areas
and extend across similar latitudes. China has all 11 genera of Pinaceae but considerably fewer
species of Pinus than the USA, while it is more diverse in other conifer genera, which mostly
explains why the two countries add up close at species rank in conifers. This is also a pointer to
the causes: the more balanced diversity across families, genera and species in China reflects a
long history of evolution without major setbacks, while North America experienced more extinc-
tions but also a few relatively recent radiations such as in Pinus and Juniperus making up for the
losses. No families but several genera are endemic to China: Cathaya, Metasequoia, Nothotsuga,
Pseudolarix and Pseudotaxus, three of these are in Pinaceae. The USA has only two endemic
conifer genera. At species rank and below the figures are again similar, with China slightly higher.
Some of that reflects different taxonomic interpretations of the observed diversity. Several taxa
with a rank of subspecies or variety in Farjon (2010a) on which this Atlas is based, were given
species rank in Debreczy & Rácz (2011) and these discrepancies are much more obvious for China
than for the USA where taxonomy has become more agreed over time. A narrower species con-
cept would give China a higher number of species but “species + infra” would remain more or
less the same figure. The total for “endemic taxa” here calculated, 124 or 55.8% is high given the
 mainland asia and japan 199

more than 20,000 km long land border of China with other Asian countries. Endemism calcula-
tions of course depend on area and with such a large country as China most species with limited
distributions within its borders will be endemic to the country. On the other hand, several such
species until recently considered endemic have now been found elsewhere, notably in Viet Nam,
such as Cunninghamia konishii, Glyptostrobus pensilis, Taiwania cryptomerioides, while conversely
Xanthocyparis vietnamensis discovered in N Viet Nam was recently found to occur in Guangxi
Province, China.

Map MAJ-35. The distribution of conifer species in mainland China and Taiwan.

The general distribution of conifer species in China shows some distinctive patterns. As with all
other maps that compile distributions of many species, density of dots on the map is influenced
by the number of species present in that area. It is therefore more informative to observe where
conifers are absent/present than to look at densities of dots; diversity distribution is better indi-
cated in map MAJ-3 on p. 174. Perhaps most striking is the absence of conifers on most of the
Tibetan Plateau, in the Kunlun Shan—Altun Shan on its N margin and in the Tarim Basin N of
these ranges. Conifers are present in the S and SE parts of the Tibetan Plateau in the drainages
of the Yarlung-Zangbo, Nu (Salween) and Lankang (Mekong) Rivers but absent from any drain-
ages northward into the plateau. These are separated by a not markedly elevated but yet signifi-
cant divide, the Nyain Qêntanglha Shan. While the S and SE parts of the Tibetan Plateau receive
monsoon rains from the E, no such influence reaches N of that divide. To the W of Gyirong the
Yarlung-Zangbo drainage is also out of reach of monsoon influence and conifers disappear from
the Tibetan side of the Himalaya along the Nepalese border. Precipitation does not increase suffi-
ciently in the Kunlun Shan—Altun Shan for conifers to thrive and conditions become only worse
in the deserts of the Tarim Basin. They reappear to the N in the Tian Shan and in mountains to
the W of the Tarim Basin (in the latter only Juniperus) but with a further gap in the N in the
Dzungarian Basin up to the base of the Altai Mountains. Near absence of conifers prevails in the
Gobi Desert with similar causes, although here Juniperus sabina var. arenaria occurs scattered
and sparsely. In the far NE of China dots on the map are thinly spread in part artificially because
of paucity of herbarium records from boreal forests. Where the Manchurian Plain becomes more
steppe-like to the W the scarcity of conifers is genuine. The North Plain only has some conifers
in the uplands of Hebei, Shandong, Henan, Hubei, Anhui and Jiangsu Provinces. This would be
200 mainland asia and japan

in part a natural scarcity as the lowlands would have been dominated by deciduous angiosperm
forest, but thousands of years of intensive agriculture make reconstruction of that situation by
pollen analysis difficult and prone to uncertainties, e.g. edaphic ‘islands’ favouring conifers may
have existed. For similar reasons the Sichuan or Red Basin has almost no conifers; the few dots
on the map are occurrences of Pinus massoniana on isolated hills. Large areas in the southern
­provinces of mainland China also lack conifers, e.g. in Hunan, Guangxi and Guangdong, but here
the distribution pattern is more diffuse as mountain ranges with natural forest are more numer-
ous. Abundance of conifer species concentrates in several areas, notably around the bend of the
Yarlung Zhangbo/Brahmaputa in Xizang [Tibet], NW Yunnan, mid-western Sichuan, NE Guangxi
and border areas with Hunan, the Anhui/Zhejiang border area, and Taiwan. In SE Xizang [Tibet],
NW Yunnan and Sichuan occur some of the richest conifer forests or mixed forests in the world,
with especially diverse representation of Cupressaceae and Pinaceae. These are montane to sub-
alpine forests in a region of high mountains and deep valleys influenced by the monsoon rains
from the Pacific Ocean. The island of Taiwan, though far to the E, is similarly diverse and the
same two families, and in one instance (Taiwania cryptomerioides) even the same species, domi-
nate the montane to subalpine forests of Taiwan. In other parts of China with high conifer diver-
sity, lying between these two conifer regions, the situation is different, with predominantly mixed
deciduous to semi-evergreen forests and a wider representation of families and genera in many
localities but with fewer species of each. It is here where the forests most resemble the earlier
Cenozoic forests which one occurred over large parts of Eurasia.
 mainland asia and japan 201

Mainland China

In this section genera and species are presented which are either endemic to mainland China or
have their major distribution there. The taxa are presented in the taxonomic sequence adopted
in Christenhusz et al. (2011) but with Cephalotaxaceae recognized at family rank and Taxaceae
in a sequence according to the latest phylogenetic results (S. Mathews, unpublished data). The
conifer species endemic to Taiwan are treated separately.

Pinaceae

All 11 genera recognized in this family are present in China and three of these are endemic:
Cathaya, Nothotsuga and Pseudolarix, while Keteleeria has its main distribution in China but
extends further S into Lao PDR and Viet Nam. Cedrus only just enters Chinese territory from the
W Himalaya (map MAJ-15 on p. 183) and the remaining six genera have wide distributions in Asia
and/or beyond in Europe and North America.

Pinus

Of the genus Pinus 24 species occur in China and of these 13 have their main distributions here,
11 in mainland China (eight endemic) and two endemic to Taiwan. We present the 11 mainland
species here, first those five that are classified in subgenus Strobus and then the six in subgenus
Pinus.

Map MAJ-36. Pinus bungeana [n=40,

m=35, h=7, 1867–1994]

Pinus bungeana has a wide range

in N-central China, but its natural
distribution is difficult to estab-
lish from herbarium collections
because this highly ornamental
species has been widely planted
for a very long time. Our map is
generally in agreement with the
one given in Ying et al. (2003)
and with the listing of provinces
under this species in Flora of China 4 (Fu et al., 1999). Other sources mention a wider distribution
but seem to include planted specimens as well. It is certainly indigenous in E Gansu, S Shaanxi,
Shanxi, Hebei, Beijing (Xi Shan), Henan, NW Hubei, N Sichuan (very rare) and Shandong (Lao
Shan). Pinus bungeana occurs in mountains, often on limestone rocks and on S-facing slopes,
scattered and mixed with P. tabuliformis and angiosperms. In the northern part of its range it is
also common on acid soils. Its altitudinal range is between 500 m and 2150 m a.s.l.; records from
lower elevations are probably not from natural occurrences, but from planted trees.
202 mainland asia and japan

Map MAJ-37. Pinus squamata [n=3, m=3,

h=3, 1991–2001]

This species is one of the rarest pines

in the world. It was discovered in
1991 in Yunnan at a locality in Qiaojia
Xian, where between 20 and 30 trees
exist on a lightly forested slope at
2200 m altitude, associated with
Pinus yunnanensis and various shrubs
and small trees in what seems to be
secondary forest. It is related to P.
bungeana (map MAJ-36 on p. 201) and
P. gerardiana (map MAJ-16 on p.
184) and all three ‘lacebark pines’
form the subsection Gerardianae
(section Quinquefolius) in the sub-
genus Strobus (Farjon, 2005b: 223).
Further searches for this pine have so far remained unsuccessful and all three herbarium col-
lections known to us were collected in the same spot. Due to its extremely limited population
P. squamata is listed as Critically Endangered (CR) on the IUCN Red List.

Map MAJ-38. Pinus armandii

var. armandii (black) [n=98,
m=92, h=13, 1888–2005]; P.
armandii var. dabeshanen-
sis (yellow) [n=7, m=7, h=5,
1956–1998]; P. armandii var.
mastersiana (red) [n=12, m=12,
h=3, 1905–2003]

Pinus armandii is widely

distributed in China and
occurs in Anhui, Chong­
qing, S Gansu, Guizhou,
Guangxi, Hainan, Henan,
Hubei, S Shaanxi, Sichuan, Yunnan, SE Xizang [Tibet], Zhejiang, Taiwan and N Myanmar [Burma].
The most common and widespread nominate variety occupies most of this range, but in a small
area in W Anhui and extreme E Hubei (Dabie Shan) it is replaced by var. dabeshanensis while in
Taiwan only var. mastersiana occurs. This is a montane species with a wide altitudinal range from
900 m to 3500 m and usually a component of mixed conifer forests. It is most commonly found
growing in forests in Yunnan and SE Xizang [Tibet] and in a wide arch surrounding the Sichuan
Basin, but with some outlying populations in S Xizang [Tibet], Guangxi, Hainan and Zhejiang
(Daxue). Such outliers could indicate that the overall distribution of P. armandii remains incom-
pletely known, but that it is rare or mostly absent in the SE of mainland China where perhaps few
mountains are extensive and high enough to support the types of cool temperate forest that this
species grows well in. The rarity of Abies in this region is a similar indicator of this.
 mainland asia and japan 203

Map MAJ-39. Pinus fenzeliana [n=56,

m=54, h=13, 1923–2009]

Pinus fenzeliana (syn. P. kwang-

tungensis) occurs in S China and N
Viet Nam, where it is locally com-
mon or abundant in the moun-
tains of Guangxi, Guangdong, S
Hunan, Hainan (island) and Ha
Giang, Son La and Hoa Binh Prov-
inces in Viet Nam. This species
occurs often on steep mountain
slopes and rocky ridges, generally
above a zone of less steep terrain
dominated by evergreen angio-
sperm trees. Its altitude range is
from (500–)700 m to 1500(–1800) m. These very steep mountainsides dry out quickly in dry sunny
weather and they sustain very little soil. Conifers are generally better adapted to cope with these
adverse conditions and the angiosperms that meet them here are smaller, deciduous and small-
leaved as opposed to the trees on the slopes below.

Pinus fenzeliana in the Nan Ling mountains of N Guangdong Province, China. Photo credit Aljos Farjon.

Map MAJ-40. Pinus wangii [n=9, m=9, h=6, 1940–1996]

Pinus wangii occurs in SE Yunnan (Malipo,

Xichou, Maquan) and possibly in Viet Nam
(Mai Chou) but the taxonomic status in Viet
Nam remains unclear (see Hiep et al., 2004). If
confirmed in Viet Nam it straddles the border
between the two countries. Its habitat is similar
to that of P. fenzeliana and the two species are
similar morphologically and closely related, but
so far P. wangii has only been found on the karst
limestone mountains (which occur in a much
wider area) whereas P. fenzeliana is also ­present
204 mainland asia and japan

on igneous rock formations. There is less difference ecologically than it would at first appear because
karst limestone is so thoroughly leached of minerals that it is almost as acidic a substrate as granite.
Pinus wangii is listed as Endangered (EN) on the IUCN Red List.

Map MAJ-41. Pinus massoniana var.

massoniana (black) [n=146, m=132,
h=17, 1803–2005]; P. massoniana var.
hainanensis (red) [n=3, m=3, h=1,
1953–1964]

Pinus massoniana is one of the

two most widespread species of
pine in China. It occurs from Sich-
uan and Yunnan to Taiwan, but
in the W of its range it does not
reach the high mountains of the
two provinces and is actually rare
in Yunnan. It is mostly a lowland
species, although it may reach
2000 m in mainland China and
2500 m in Taiwan, distributed in
warm temperate regions but with more continental conditions (colder winters) in the interior of
China. Readily invading abandoned fields, especially in rocky terrain on acidic soil, it is a species
that has taken advantage of forest degradation and can even occur on hills in the midst of exten-
sive rice plantations, e.g. in the Sichuan Basin. On Hainan Island a distinct variety P. massoniana
var. hainanensis is recognized; it is listed as Critically Endangered (CR) on the IUCN Red List.

Map MAJ-42. Pinus tabuliformis

[n=111, m=108, h=12, 1875–2008]

Pinus tabuliformis is another wide-

spread pine of China. It has two
rather obscure varieties which
we do not show on the map for
that reason and because her-
barium specimen data found are
incomplete to show their distri-
butions. This species has a much
more northerly distribution than
P. massoniana with little overlap-
ping of ranges; the two species
meet in S Shaanxi, Chongqing, W Hubei and a few other places further E. In Sichuan P. tabuli-
formis occurs in the higher mountains W and S of the range of P. massoniana and there is one
record from NW Yunnan (Heqing, coll. H. von Handel-Mazzetti No. 8734 in 1916). In the NE of its
range P. tabuliformis extends to Jilin Province and to North Korea. The altitude range is greater
than that of P. massoniana and it can occur to 3800 m in Sichuan. Its ecology is similar, but it is
adapted to lower winter temperatures, hence it more northerly distribution and capacity to reach
greater altitude in the W mountains of S Gansu and Sichuan.
 mainland asia and japan 205

Map MAJ-43. Pinus yunnanensis var.

yunnanensis (black) [n=71, m=65,
h=12, 1874–2005]; P. yunnanensis
var. pygmaea (red) [n=11, m=10, h=3,
1914–1986]

Pinus yunnanensis is distributed

mainly in Yunnan and Sichuan
but extends westwards into SE
Xizang [Tibet] and occurs in a
few localities in Guizhou and
W Guangxi. A shrubby form,
P. yunnanensis var. pygmaea, is
found scattered within the main
range of the species and usually
at higher altitudes. The altitude
range of the species is from 400 m
to 3300 m, forming extensive forests on S-facing slopes in river valleys. It is most commonly gregari-
ous in pure stands, but can be found growing with other pines, e.g. P. armandii, P. tabuliformis and
P. kesiya, or with Keteleeria evelyniana, as well as with angiosperm trees and shrubs. Near Weixi
in W Yunnan it has been found associated with Torreya fargesii var. yunnanensis (Debreczy &
Rácz, 2011: 421). On rocky slopes P. yunnanensis is often the only pine in open scrubland and above
2200 m the krummholz variety pygmaea may occur among shrubby oaks, cotoneasters and other
xerophytes.

Map MAJ-44. Pinus densata [n=60,

m=56, h=9, 1904–2003]

Pinus densata occurs in the moun-

tains of W Sichuan, NW Yunnan
and SE Xizang [Tibet]. In Ying
et al. (2003: 74) the map for this
species is similar to ours (note that
the SW border of China includes
Arunachal Pradesh, claimed by
China but under Indian admin-
istration) but it has a dot in E
Qinghai which is not confirmed
by our data. The species is also
reported from S Qinghai but again no herbarium specimens were found. Pinus densata appears
to be endemic to China in whichever way one draws the border with India, but some localities
are on or near the actual border and it would be no surprise if it was found in Arunachal Pradesh
now that access to the region has become easier. It is a high altitude species occurring from 2600
m to 4200 m, at the higher elevations forming pure stands but lower down mixed with other
pines such as P. armandii and P. yunnanensis. Its range is mainly further W than P. tabuliformis;
its altitude range is above that of P. yunnanensis. Several authors have suggested that P. densata
may be of hybrid origin with these species, or a third unknown species, as parents, based on mor-
phological as well as molecular evidence (Liu et al. 2003). If so, P. densata has apparently spread
to areas and altitudes not now occupied by its parental species, indicating a long time lapse since
hybridization began.
206 mainland asia and japan

Map MAJ-45. Pinus henryi [n=34, m=31,

h=7, 1888–1998]

Pinus henryi has a limited distribu-

tion in Chongqing (formerly E part of
Sichuan), W Hubei, Hunan, S Shaanxi
and N Sichuan. It is most common
in W Hubei and in the Daba Shan in
Chongqing where it occurs at middle
elevations from 1100 m to 2000 m in
usually dry locations in more or less
open woodland or on rocky slopes
and outcrops. This species is closely
related to P. tabuliformis (map MAJ-42
on p. 204) and is treated by some authors as a variety of that species. Their ranges do partly
overlap especially in the Daba Shan, but in the most W part of Hubei we only have herbarium
records of P. henryi.

Map MAJ-46. Pinus hwangshanensis

[n=53, m=53, h=7, 1924–2011]

Pinus hwangshanensis occurs in SE

China from Guangxi in the W to Zhe-
jiang in the E. In Chinese accounts
(e.g. Flora of China 4; Fu et al., 1999)
this species is synonymous with
P. taiwanensis but Farjon (2001; 2005;
2010a) and Debreczy & Rácz (2011)
treat these two as distinct species,
which is followed here. This species
usually occurs on steep rocky ridges
and can grow even from crevices in
bare rock. At these extreme localities
it evades competition from nearly all other trees and often the crags and summits are adorned
with the picturesque flat-topped and contorted pines so often accurately drawn in traditional
Chinese landscape paintings. Although some other Chinese species of Pinus can occupy steep
ridges, e.g. P. fenzeliana, no other species matches P. hwangshanensis in its capacity to establish
and grow, no doubt slowly, in such extreme situations.

Pinus hwangshanensis on Bijia Shan, Jinggangshan, Luoxiao Range, Jiangxi Province, China. Photo credit
Aljos Farjon.
 mainland asia and japan 207

Cathaya

Map MAJ-47. Cathaya argyrophylla

[n=28, m=27, h=9, 1938–1996]

Cathaya argyrophylla is the only

extant species in this genus,
endemic to China. It occurs in
a limited number of mostly dis-
junct locations in S Chongqing
(formerly part of Sichuan), N Gui-
zhou, NE Guangxi and E Guangxi
and S Hunan. Some consist of
subpopulations in a number of
nearby localities and some are
only a small population in a
single locality. Its much wider
distribution in the Cenozoic is
confirmed by a relatively abundant fossil record, especially from Central Europe where Cathaya
(possibly the same species) occurred until the beginning of the Pleistocene. Fossils are also known
from France, Greece, Georgia and Siberia, while pollen ascribed to this genus has been found in
the Eocene of the Canadian Arctic. Cathaya argyrophylla occurs in medium high mountains at
altitudes between 900 m and 1900 m, usually with other conifers e.g. Pinus fenze­liana, Tsuga chin-
ensis and Nothotsuga longibracteata, in localities where evergreen angiosperm forest gives way to
semi-deciduous forest on steeper slopes and on crags above ca. 1000 m. Although this species is
protected and occurs in several protected areas, its small area of occupancy (AOO) is sufficient
reason to list it as Vulnerable (VU) on the IUCN Red List.

Picea

The genus Picea is extraordinarily diverse in China; of a total of 38 species in this circumpolar
genus 19 occur in China. Ten species are endemic to mainland China and one species is endemic
to Taiwan. Some species have wider occurrence outside China and are treated in other sections of
this chapter. This leaves 12 species to be treated here and one in the section on Taiwan endemics.
The distribution of the genus Picea in China is interesting because of its absence in the mainland
E and S and isolated occurrence in Taiwan. Picea is a genus of cool and cold climates and spruces
migrate northwards and upwards when the climate warms. In E and S China no mountain ranges
are high enough for any species of Picea; such mountains do exist in Taiwan only and W of the
island the first mountain ranges with an average altitude >2000 m are in W Hubei. N of the
40th parallel forests with Picea begin to appear at much lower altitudes. The species diversity is
concentrated in the high mountains of S Gansu, W Sichuan, NW Yunnan and to a lesser degree
in SE Xizang [Tibet]. Here repeated retreat to refugia during glacial maxima and return during
interglacial times in the Pleistocene must have caused isolation of populations, hybridization and
speciation, leading to the present taxonomic complexity and variation. In the N species diversity
drops sharply; forests usually have only one, sometimes two species of Picea spreading over large
territories.
208 mainland asia and japan

Map MAJ-48. The distribution of the genus Picea in mainland China and Taiwan.

Map MAJ-49. Picea asperata [n=52,

m=48, h=11, 1903–1996]

Picea asperata is a forest forming

species distributed widely in the
high mountains of S Gansu, SW
Shaanxi and Sichuan, with some
outlying populations to the N in
the Helan Shan in Ningxia and
to the W in E Qinghai Province.
Besides the most common nomi-
nal form two varieties are recog-
nized, but not shown on the map
as few specimens are available or
correctly identified and they do
not occur in separate areas. The
altitude ranges from 1500 m to
3800(–4100) m, usually above 2400
m in Sichuan. In drier regions in
the N of its range P. asperata forms nearly pure spruce forests on N-facing slopes, in Sichuan
other species of conifer commonly associate with it, e.g. Abies chensiensis, A. fargesii and Pinus
tabuliformis. Extensive logging has reduced old growth spruce forests in the region dramatically,
persuading the central government of China to issue a general logging ban in these forests. This
species is an important timber tree and the subject of replanting schemes within its natural range.
Only six of the records in the Conifer Database used to produce this map date from 1980 or later
and it is likely that all refer to old growth forests.
 mainland asia and japan 209

Map MAJ-50. Picea aurantiaca [n=9, m=9, h=7, 1903–1989]

Picea aurantiaca is closely related to P. asperata (map

MAJ-49 on p. 208) and sometimes treated as another
of its several varieties. It has a very limited distribu-
tion confined to the part of the Daxue Shan (Zheduo
Shan) near and NW of Kangding (“Tachien-lu” on
older herbarium labels). It occurs mostly in limestone
formations at altitudes between 2600 m and 4000 m in
subalpine conifer forest with e.g. Picea likiangensis var.
rubescens, Abies squamata, A. chensiensis, A. recurvata, Tsuga chinensis and locally Larix potaninii.
Betula spp. are the common broad-leaved trees, while Pinus spp. occur mostly after disturbances
and at the lower elevations. Picea aurantiaca is listed as Endangered (EN) on the IUCN Red List
because of intensive logging in the past.

Map MAJ-51. Picea retroflexa [n=15, m=15, h=10, 1903–1984]

Picea retroflexa is another species closely related

to P. asperata. Although mostly confined to the
mountains of W Sichuan, it has been collected
in Banma Xian in S Qinghai and in Jone Xian,
S Gansu. It is most common in mountains between
the Yalong River and the Xianshui River NW of
Kangding, with summits rising above 5000 m. Picea
retroflexa is a species of subalpine conifer forests,
found at altitudes between 3000 m and 4700 m (the
latter altitude is reached E of Dawu, Sichuan). At the
highest elevations it grows either pure or mixed with
Abies squamata. Picea retroflexa is listed as Endan-
gered (EN) on the IUCN Red List due to decline
caused by logging.

Map MAJ-52. Picea brachytyla var.

brachytyla (black) [n=34, m=33, h=12,
1900–2004]; P. brachytyla var. com-
planata (red) [n=36, m=36, h=10,
1903–2000]

The two varieties of Picea brachy-

tyla occur broadly in the same
region of China, but var. com-
planata becomes the more
common variety in the W part, i.e. NW Yunnan and SE Xizang [Tibet], while in Sichuan both
varieties occur more or less with equal frequency. In the N and E of the species range (S Qing-
hai, N Sichuan, S. Gansu, S. Shaanxi and W Hubei) only var. brachytyla has been collected. The
species (presumably P. brachytyla var. complanata) is recorded from Arunachal Pradesh in NE
India and N Myanmar [Burma] and possibly Bhutan, but no herbarium collections from these
areas were found. Recent surveys have collected the species frequently in the Gaoligong Shan
in NW Yunnan close to the Myanmar border and it almost certainly occurs on the other side
of the divide, but access to the most northern part of this country remains very difficult and
no collections are likely to be made soon. Picea brachytyla is a high montane to subalpine spe-
cies and a constituent of mixed conifer forests at altitudes between 1300 m and 3800(–4000) m.
Picea brachytyla is listed as Vulnerable (VU) on the IUCN Red List.
210 mainland asia and japan

Picea brachytyla (darker trees) in mixed conifer forest at Bitahai Lake, Zhongdian Xian, Yunnan, China.
Photo credit Aljos Farjon.

Map MAJ-53. Picea purpurea [n=38, m=37,

h=6, 1903–1993]

Picea purpurea has a similar distribution as

P. asperata (map MAJ-49 on p. 208) and
occurs in S Gansu, extreme E Qinghai,
W-central and SW Sichuan and extreme
NW Yunnan. Like P. asperata it occurs
in mixed or sometimes pure conifer for-
ests between 2600 m and 3800 m, most
commonly on N-facing slopes. Near
Songpan in NE Sichuan the two spe-
cies occur together in high montane
spruce forest. On very steep rock slopes
Picea purpurea is often the only spruce, and in such situations junipers may be the only other
conifers present.

Map MAJ-54. Picea likiangensis [n=95, m=93,

h=20, 1884–2004]

Picea likiangensis is a variable species

in which four or five varieties are recog-
nized (Farjon, 1990, 2001, 2010a); one of
these was raised to species rank which
is here accepted (see P. linzhiensis map.
MAJ-55 on p. 211). The distribution of
these varieties is largely sympatric (var.
montigena is limited to Sichuan) and
they are not separately shown on the
map. Picea likiangensis occurs in the high mountains of S. Qinghai, W Sichuan, NW Yunnan and
SE Xizang [Tibet]. This too is a subalpine spruce at altitudes between 2700 m and 4100 m where
it occurs with e.g. P. brachytyla, P. purpurea, P. asperata, Abies squamata at the highest altitudes
and A. forrestii, Larix potaninii and Tsuga spp. somewhat lower down the slopes. The species is
listed as Vulnerable (VU) on the IUCN Red List.
 mainland asia and japan 211

Map MAJ-55. Picea linzhiensis [n=13,

m=13, h=4, 1936–2002]

Picea linzhiensis is closely related

to P. likiangensis and by several
authors (e.g. Debreczy & Rácz,
2011: 387) treated as a variety of it.
Its distribution partly coincides
with the range of P. likiangensis
in the far NW of Yunnan (Dêqên
Prefecture) and SE Xizang [Tibet] but it mostly extends further W in Xizang than that species and
has been collected from Bomi Xian as far W as Yatung. Although the type specimen was collected
from Linzhi on the lower Yarlung Zangbo River, most of these localities are near the borders with
India (Arunachal Pradesh) and Bhutan but no records are as yet known from these countries. We
also have not found herbarium records from SW Sichuan; a careful search in Chinese herbaria
will perhaps reveal these but such detailed investigation fell beyond the scope of the work on the
Atlas. In SE Xizang this species forms almost pure forests between 3000 m and 3800 m, usually
well above a mixed coniferous forest belt in which Picea spinulosa is the dominant spruce. At
around 3000 m it occurs also mixed with Pinus armandii, while at its upper limit it grows with
Larix sp. and Abies spp., the firs ultimately replacing the spruces above 3600–3800 m a.s.l.

Map MAJ-56. Picea crassifolia [n=50,

m=41, h=5, 1911–1994]

Picea crassifolia is widespread

in the mountains of E Qinghai,
Gansu and Ningxia. In Nei Mon-
gol (Inner Mongolia) it is rare and
limited to a few isolated locations
(Daqing Shan, Lang Shan, Xilin-
hot). In the Helan Shan it forms
forests on either side of the moun-
tains (Ningxia on the E slopes, Nei
Mongol on the W slopes) and grows on limestone. It is mostly confined to N-facing slopes, above
steppe or desert, at elevations between 1600 m and 3800 m a.s.l. The climate is cold continental
and dry, with most of the precipitation falling as snow. Picea crassifolia forms mostly pure forests,
here and there with Betula albosinensis and groves of Populus tremula. At lower altitude it can
occur mixed with Pinus tabuliformis (mostly in S Gansu) and Juniperus rigida.

Map MAJ-57. Picea meyeri [n=36, m=33, h=9,

1913–1976]

Picea meyeri is distributed across the moun-

tains that separate the Northern Plain from the
Gobi Desert, occurring in N Shanxi, N Hebei
and parts of Nei Mongol (Inner Mongolia). In
some regions of these mountains it occurs near
the Great Wall, elsewhere as in Shanxi mostly
to the S in e.g. the Heng Shan and Wutai Shan.
It is reported from Shaanxi Province but no
herbarium collections were found; presumably
212 mainland asia and japan

it may occur just inside the border with Shanxi. Picea meyeri is a high montane to subalpine spe-
cies, occurring at elevations between 1600 m and 2700 m, often restricted to the N-facing slopes
of the mountains. It grows in pure stands or mixed with Picea wilsonii, Abies nephrolepis and Larix
gmelinii var. principis-rupprechtii, the latter at elevations above 2100 m on Xiaowutai Shan.

Map MAJ-58. Picea wilsonii [n=64,

m=59, h=12, 1901–2000]

Although Picea wilsonii occurs in

the same northern mountains as
P. meyeri (map MAJ-57 on p. 211)
it extends much further W and its
main range is in S Gansu, N Sich-
uan, Shaanxi, Chong­qing (Daba
Shan) and W Hubei. An outlying
population occurs in the SW of
Sichuan near the village Hetaowan
(Muli Zangzu Zizhixian). As can be
expected with such an extensive
distribution the altitude range is
considerable, with a lowest record at 1400 m and to 3000 m a.s.l. in Sichuan. In the montane
sub-boreal coniferous forests of N Sichuan and S Gansu it is usually growing with Abies fargesii,
A. recurvata, Picea asperata, P. purpurea and Pinus tabuliformis. In the NE of its range Picea wil-
sonii occurs also with P. meyeri and Abies nephrolepis or in pure stands. At its lowest altitude
angiosperms such as Betula, Acer, Ulmus and Salix are important constituents of mixed deciduous
forest here.

Map MAJ-59. Picea neoveitchii [n=14,

m=12, h=6, 1901–1982]

Picea neoveitchii is extremely rare

and known only from scattered and
very small populations, in some cases
only a single mature tree. This spe-
cies has been collected in S Gansu,
S Shaanxi, W Hubei and SW Henan.
In the Western herbaria we only have
E. H. ­Wilson’s collections, including
the type, from W Hubei at the begin-
ning of the twentieth century; all
other and later specimens are in Chinese herbaria. Some of the populations mapped here may no
longer exist because they occur in logged-over mountains or valleys. Some other localities have
trees of uncertain identity that may be of hybrid origin. Picea neoveitchii occurs in medium high
mountains, at elevations between 1200 m and 2250 m, often on N-facing, steep, rocky slopes. With
only a few of the known populations in forest reserves, this species appears to be still declining
and is listed as Critically Endangered (CR) on the IUCN Red List.
 mainland asia and japan 213

Map MAJ-60. Picea koraiensis var. koraiensis

(black) [n=17, m=15, h=7, 1897–1993]; P. koraiensis
var. pungsanensis (red) [n=1, m=1, h=1, 1940]

Picea koraiensis has a distribution in the far-

thest NE provinces Jilin and Heilongjiang in
China and also occurs in North Korea and in
Primorye in the Russian Far East. The few her-
barium collections available from North Korea
are from 1917 (E. H. Wilson, V. Komarov) and
1935 and 1940 (var. pungsanensis); this species
is undoubtedly more widespread there. Some
named localities in North Korea could not be
found for the map due to non-traceable name
changes. Picea koraiensis var. pungsanensis is
only known from its type locality, Mt. Pung-
san (=Puksubaek-san) but the whereabouts
(in Japan) of the type specimen is uncertain. This species occurs in mixed angiosperm-conifer
forests which form a transitional zone between the deciduous forests to the S and the evergreen
boreal conifer forests to the N and NE.

Pseudotsuga

Map MAJ-61. Pseudotsuga sinensis

var. sinensis (black) [n=58, m=51, h=18,
1906–2004]; P. sinensis var. brevifolia
(red) [n=16, m=16, h=5, 1956–2004];
P. sinensis var. gaussenii (yellow)
[n=11, m=11, h=8, 1924–1991]

The genus Pseudotsuga has just

one variable species in main-
land China, Pseudotsuga sinen-
sis, which also occurs in Taiwan.
It is here mapped with its three
varieties, of which P. sinensis var.
sinensis is the most widespread and occurs from extreme SE Xizang [Tibet] and NW Yunnan
to Taiwan. [At both extremities the populations have been recognized as distinct species.] The
distribution of this variety is mainly concentrated in SW-central China, i.e. Sichuan, Yunnan,
Chongqing, W Hubei, Guizhou and W Guangxi. One or two localities occur in N Jiangxi and it is
present in the central mountains of Taiwan. From Guangxi it crosses the border into N Viet Nam.
Pseudotsuga sinensis var. brevifolia has a more limited distribution and occurs in S Guizhou, in
Guangxi and in N Viet Nam. It may also be present in the extreme SE of Yunnan, as some of the
Vietnamese populations are close to the border. Pseudotsuga sinensis var. gaussenii is distributed
in S Anhui, NE Jiangxi (Dexing) and Zhejiang, but was also collected in N Guizhou. The distribu-
tion of the latter variety is probably incompletely known, in part due to identification problems
with sterile (no seed cones) dried herbarium specimens. Pseudotsuga sinensis occurs at various
altitudes from 600 m to 3300 m, with the highest in NW Yunnan. This conifer is a relatively rare
constituent of mixed mesophytic forests, where sometimes angiosperms (deciduous or evergreen)
and in other places conifers dominate. Pseudotsuga sinensis is listed as Vulnerable (VU) on the
IUCN Red List because of deforestation in large parts of its range.
214 mainland asia and japan

Pseudotsuga sinensis in Taroko National Park, Central Mountains, Taiwan. Photo credit Aljos Farjon.

Larix

There are five species of Larix in China: L. gmelinii, L. griffithii, L. mastersiana, L. potaninii and
L. sibirica. Of these, only L. mastersiana is endemic to China, but several varieties recognized
under these species are also limited to China. Species with wider distributions outside China are
treated in other sections of this chapter, leaving besides L. mastersiana only L. potaninii to be
treated here, but with the exception of L. potaninii var. himalaica which is mapped in the section
on the Himalayas (map MAJ-22 on p. 188).

Map MAJ-62. Larix mastersiana [n=13, m=11, h=8,

1908–1985]

Larix mastersiana is a rare species with a lim-

ited distribution in Sichuan, mainly in the
mountains in the central part of the province
W of the Sichuan Basin. The mountains on
either side of the Min River Valley have sev-
eral populations, but more isolated localities
with this species are known to the SW in Baox-
ing Xian and Li Xian, as well as in the Dadu
River drainage N of Kangding. Two localities
in N Sichuan are wide apart, the most western
one was collected by F. Kingdon-Ward in 1935,
the other by a “Sichuan Vegetation Survey Team” in 1985. The paucity of herbarium collections
and the time intervals attest to the rarity of this species. Some localities based on collections
made in the early decades of the twentieth century may no longer have any trees left; a recent
survey found only the following three locations: Min River drainage, Dadu River drainage, and
“upstream of Qingyijiang” (State Forestry Bureau, 2009) an unintelligble locality: the river Qingyi
Jiang discharges into the Min River near Leshan S of Chengdu and originates in S Sichuan, far
from any known populations. The altitude range of this species is 2000–3500 m and it occurs
mostly in wet mixed conifer forests with Abies fargesii, A. forrestii, Picea purpurea and in the N
P. wilsoniana. Larix mastersiana is listed as Endangered (EN) on the IUCN Red List.
 mainland asia and japan 215

Map MAJ-63. Larix potaninii var.

potaninii (black) [n=62, m=62, h=10,
1891–2001]; L. potaninii var. chinen-
sis (red) [n=6, m=5, h=3, 1894–1958];
L. potaninii var. macrocarpa (yellow)
[n=14, m=14, h=5, 1914–1995]

Larix potaninii is widespread in

the Sino-Himalayan mountain
system, which comprises the
eastern part of the Himalaya and
the adjacent but more N-S ori-
ented ranges in E Xizang [Tibet],
Sichuan and Yunnan. The three
varieties mapped here (for var. himalaica see map MAJ-22 on p. 188) are endemic to China as far
as is known, but L. potaninii var. potaninii may also occur in India (Arunachal Pradesh) and N
Myanmar [Burma] given its known distribution in SE Xizang and NW Yunnan. This is the most
common and widespread form and it occurs as far N as S Gansu and S Shaanxi. Larix potaninii var.
macrocarpa occurs with it in SW Sichuan and NW Yunnan, but it is a somewhat ill-defined taxon
based on larger seed cone size and so sterile herbarium specimens cannot easily be identified and
this variety may be under-represented in the records. Larix potaninii var. chinensis has a limited
known distribution in the Qin Ling (mountain range) in Shaanxi province. All varieties are high
montane to subalpine trees occurring from 2350 m to 4300 m altitude, forming usually scattered
stands among other conifers or with deciduous angiosperms at lower elevations. As light requir-
ing trees these larches often benefit from avalanches that have cleared away ‘dark’ conifers such
as Abies and Picea and their stands may mark old avalanche or rockfall paths.

Pseudolarix

Map MAJ-64. Pseudolarix amabilis [n=28,

m=25, h=8, 1907–1974]

The natural distribution of Pseudolarix

amabilis, the only species in this genus
and an endemic of China, is difficult
to establish. This tree, with its colour-
ful autumn foliage, has been planted in
many places where it was not originally
present; in more recent times forestry
has taken an interest in it as well. Old
growth forest with this species is cer-
tainly rare now. In Atlas of the Gymno-
sperms of China (Ying et al., 2003), a map
with dots (based on herbarium specimen data from Chinese herbaria) is produced which gives a
much wider range than here presented. A map given in China Plant Red Data Book 1 (Fu & Jin,
1992) is similarly optimistic. A check by Dr Xiang Qiao-ping in the Beijing Herbarium (PE) revealed
that most of these specimens were collected from planted trees or trees in villages and towns
(e.g. near temples) that did not occur in the wild. This problem is acknowledged in Flora of China
4 (Fu et al., 1999). The map presented here gives localities believed to be old growth forest with
this species, which is deciduous or semi-evergreen angiosperm forest in which P. amabilis occupies
216 mainland asia and japan

steep slopes, rocky outcrops and ridges where competition is less intense. Its altitudinal range is
from 180 m to 1200 m, so it is a component of lowland forest in a warm temperate, humid climate
which can experience occasional cold winter frost at higher altitudes. On the famous mountain
Tianmu Shan in Zhejiang it occurs on sandstone ridges and steep slopes together with Ginkgo
biloba and Pinus hwangshanensis at ca. 1000 m a.s.l. in deciduous forest. Although this and other
localities are protected, there is still the risk of (illegal) felling of trees, which can be tall and straight
and produce good timber. Pseudolarix amabilis is listed as Vulnerable (VU) on the IUCN Red List.
From an evolutionary perspective this genus is of much interest and during most of the Cenozoic
it was widely distributed in the Northern Hemisphere, in Eurasia as well as in North America.

Tsuga

Three species of Tsuga occur in China, one of these, T. dumosa, is primarily a Himalayan species
and has been treated in that section of this chapter (map MAJ-23 on p. 189).

Map MAJ-65. Tsuga chinensis [n=140, m=130,

h=20, 1888–2011]

Tsuga chinensis has a very wide distribu-

tion in mainland China; it also occurs in
two provinces (Ha Giang and Cao Bang)
near the Chinese border in Viet Nam, and
in Taiwan. Three varieties are recognized
but herbarium data are insufficient to give
their distribution so we present a map
of the species. It is not certain whether
T. chinensis occurs in SE Xizang [Tibet]
as stated in Flora of China 4: 4 (Fu et al.,
1999) and Farjon (2010a) because no herbarium specimens to confirm this were found for the map
in Atlas of the Gymnosperms of China (Ying et al., 2003) or for this Atlas. Probably only T. dumosa
occurs in Xizang (map MAJ-23 on p. 189). Tsuga chinensis and its varieties occur at altitudes of
(600–)1200–3200(–3500) m in a number of different forest types, from temperate deciduous angio-
sperm forest in the E to montane-subalpine conifer forest in the W. In the lower mountains of E and
S China it is much less widespread than in the high mountains of Sichuan and Yunnan, but where
present it can be one of the commonest conifers. In these warmer regions it occurs on acidic rocks
such as granite or quartzite/sandstone as well as on limestone, in the S of China and in Viet Nam
typically on karst mountains; more gentle slopes, if still forested, are dominated by angiosperms.

Tsuga chinensis (left) with Fokienia hodginsii (right) on a steep slope of Bijia Shan, Jinggangshan, Luoxiao
Range, Jiangxi Province, China. Photo credit Aljos Farjon.
 mainland asia and japan 217

Map MAJ-66. Tsuga forrestii [n=17, m=17, h=7,

1913–2000]

Tsuga forrestii has a much more limited distribu-

tion than T. chinensis and is with certainty known
from W-central and SW Sichuan and NW Yunnan.
Ying et al. (2003) give a map with dots also in NE
Guizhou and W Hubei, but the latter are almost
certainly incorrect identifications of T. chinensis.
According to Flora of China 4 (Fu et al., 1999) and
Farjon (2010a) it does occur in the Fanjing Shan,
Jiangkou Xian in NE Guizhou (ca. 27° 48’ N 108° 46’ E) but we have no herbarium specimen records
from there in the Conifer Database. This would be an easternmost outlying population of this species.
Tsuga forrestii is a high mountain species occurring between 2000 m and 3500 m a.s.l. It is a constituent
of high montane conifer forests, where it is mixed with Abies spp., Picea spp., Larix potaninii, occasion-
ally Pseudotsuga sinensis, Cephalotaxus fortunei, and angiosperm trees, e.g. Betula albosinensis, Acer
spp., Sorbus spp. Quercus spp., and Magnolia spp. Tsuga forrestii remains in most places a minor com-
ponent of the forest (Wang, 1961). Tsuga forrestii is listed as Vulnerable (VU) on the IUCN Red List.

Nothotsuga

Map MAJ-67. Nothotsuga longibrac-

teata [n=52, m=51, h=11, 1928–2009]

The genus Nothotsuga has a

single species, N. longibracteata
which is endemic to China. It has
a wide but mostly scattered dis-
tribution in SE China and occurs
in NE Guizhou, Hunan, Guangxi,
N Guangdong, SE Jiangxi and
Fujian. It is rare to locally com-
mon in mountains from ca. 300 m
to 2300 m in evergreen sclerophyl-
lous angiosperm forest (Castanop-
sis, Lithocarpus, Quercus) at lower
elevation and in mixed angiosperm/conifer forest at higher elevation. In the latter forest type
associated conifers are e.g. Tsuga chinensis, Pinus fenzeliana, P. massoniana and Torreya grandis.
Logging has depleted mature tree stands in many of the more accessible localities lower down the
slopes, so that now most large trees are found above the steepest crags on mountain ridges from
where transport of logs would be difficult. A number of populations are in protected areas and a
general ban on logging old growth forest also helps towards the conservation of this interesting
species. It is assessed as Near Threatened (NT) on the IUCN Red List, i.e. at present not meeting
criteria for a threatened category (VU, EN or CR) but perhaps on its way to that status.
218 mainland asia and japan

Nothotsuga longibracteata in the Nan Ling mountains, S Hunan, China. Photo credit Aljos Farjon.

Keteleeria

The genus Keteleeria has three species and is primarily distributed in China, including Taiwan, but
one species extends S into Lao PDR and Viet Nam. All three are somewhat variable in different
locations, having led to the recognition of several species by some Chinese and European authors.
The distribution of the genus shows some regions where it is common, such as S Sichuan and
N-central Yunnan, W Hubei and N Guangxi and adjacent parts of Guizhou and SW Hunan, but
also areas of not easily explicable absence, such as most of Hunan (see also near absence of Taxus
in that province, maps MAJ-123–124 on p. 251) and Jiangxi, N Guangdong and much of Fujiang and
Zhejiang. In Taiwan it is mostly limited to the far N of the island, even though this area is densely
populated with the capital city in its centre. Absence in some areas of SE China may be partly due
to large scale deforestation, as the genus occurs in lowland forests as well as at higher montane
elevations. Particularly in the E of China native forests are now often restricted to isolated rem-
nants associated with Buddhist monasteries or shrines, which often incorporated a forested hill
or mountain slope in the sanctuary and thereby effectively practised nature conservation.

Map MAJ-68. The distribution of the genus

Keteleeria.
 mainland asia and japan 219

Map MAJ-69. Keteleeria davidiana

var. davidiana (black) [n=61, m=58,
h=15, 1869–1997]; K. davidiana var.
formosana (red) [n=15, m=9, h=8,
1902–1997]

Keteleeria davidiana var. davidi-

ana is widely distributed in
S-central China and occurs in
Yunnan (uncommon), Sichuan,
extreme SE Gansu, Chongqing,
S Shaanxi, W Hubei, Guizhou,
Hunan and Guangxi. It has also
been found in Bac Can Province, N Viet Nam. In its westernmost part of the range it is less fre-
quent, being replaced by K. evelyniana. Keteleeria davidiana var. formosana is endemic to Taiwan.
Only small populations of this variety occur, concentrated in the far N of the island and only two
in the S in Hualien and Pingtung Counties, both isolated. This variety is in danger of extinction
due to encroaching development in the N and the expansion of palm (Areca catechu) plantations
in the S and is listed as Endangered (EN) on the IUCN Red List. Keteleeria davidiana occurs from
hills to low mountains at elevations of (300–)600–1000(–1300) m in a warm-temperate to sub-
tropical, moist climate. It is a constituent of the mixed mesophytic forest formation (Wang, 1961),
together with many genera and species of broad-leaved deciduous trees, and some other gym-
nosperms, such as Pinus massoniana, P. bungeana, Cunninghamia lanceolata, Cupressus funebris,
Torreya grandis, and Podocarpus nakaii (Taiwan). It also occupies the evergreen broad-leaved
forest formation (Guangxi, Guizhou, Taiwan), with numerous (sclerophyllous) evergreen tree spe-
cies and Pinus spp. It rarely forms pure stands. Keteleeria survives coppicing and, like many spe-
cies of Pinus, appears to act as a pioneer in secondary vegetation.

Map MAJ-70. Keteleeria fortunei [n=42, m=41,

h=12, 1852–2002]

Keteleeria fortunei occurs in S China in

S Guizhou, SE Yunnan, Guangxi, S Hunan,
Guangdong, Hong Kong, S Jiangxi, Fujian
and S Zheijiang. The altitude range is
between 350 m and 1400 m and it is com-
monly found in mixed evergreen angio-
sperm-dominated forest where it can
compete with other trees to reach the
canopy and become sometimes domi-
nant, as in a protected forest near the Yongquan temple close to the city of Fuzhou in Fujian. Much
of this lowland forest has been cleared a long time ago and therefore the distribution of K. fortunei
has become much more fragmented than it must have been in the past. Although this species is a
co-dominant tree in primary forest, it can establish itself also in secondary forest and does so where
seed sources are not too distant.
220 mainland asia and japan

Map MAJ-71. Keteleeria evelyniana [n=67, m=62, h=16,

1898–2007]

Keteleeria evelyniana is distributed from SW Sichuan

through much of central Yunnan and, more disjunct,
in SE Yunnan, Hainan, N Viet Nam and adjacent parts
of Lao PDR and in the Lang-Bian Massif in Lam Dong
and Ninh Thuan Provinces in S Viet Nam. In Yunnan
this species occurs in the high mountains up to 2700–
3000 m; in Viet Nam it occurs at 1900 m on Bi Doup
Mountain in Lam Dong Province. Its lowest altitude
recorded on herbarium labels is 750 m in N Viet Nam.
It may have been a constituent of lowland subtropical
evergreen forest but in many places in the hills and
lower mountains of Yunnan these forests have been
logged and turned into scrub in which K. evelyniana
maintains itself as a shrub or stunted tree. Keteleeria
evelyniana is listed as Vulnerable (VU) on the IUCN
Red List mainly due to widespread deforestation.

Abies

The genus Abies is well represented in China and Taiwan with 16 species, one of these is endemic
to Taiwan and 10 are endemic to mainland China. Abies densa (map MAJ-24 on p. 189) and A. spect-
abilis (map MAJ-25 on p. 190) are Himalayan species treated in that section of the chapter; A sibirica
(map MAJ-13 on p. 181 occurs mostly in Siberia. The Taiwan endemic, A. kawakamii (map MAJ-129
on p. 253) is given a map and discussion separately with other Taiwan endemic conifers.

Map MAJ-72. The distribution of the

genus Abies in mainland China and
Taiwan.

The distribution of the species of

Abies in China is concentrated in a
relatively limited area which coin-
cides with the high mountains
of S Gansu, Sichuan, NW Yun-
nan and SE Xizang [Tibet], with
an extension into S Shaanxi and
W Hubei to the E and along the
Himalaya to the W. In other parts
of China fewer species occur and these populations are separated by greater distances except in
the NE where Abies nephrolepis occurs continuously with its distribution in the Russian Far East.
Similarly, the occurrence of A. sibirica in the far N of Xinjiang is just an extension of that species
into China in the Altai Mountains. The single species in Taiwan is also disjunct, while four spe-
cies in SE China are extremely restricted in one or a few tiny populations. The taxonomy of the
high mountain species, as in Picea, is complicated with several polymorphic species, continuous
variation and possibly hybridization, indicating recent or indeed ongoing speciation; this is
reflected in difficulties encountered with phylogeny reconstruction for the genus (Xiang et al.,
2004, 2009).
 mainland asia and japan 221

Map MAJ-73. Abies chensiensis ssp. chen-

siensis (black) [n=15, m=13, h=4, 1872–1997];
A. chensiensis ssp. salouenensis (red) [n=18,
m=17, h=8, 1913–2003]; A. chensiensis ssp.
yulongxueshanensis (yellow) [n=4, m=4,
h=5, 1923–1994]

This species has a disjunct distribution,

with Abies chensiensis ssp. chensiensis in
the N (S Gansu, S Shaanxi, W Hubei, SW
Henan) separated from the two other
subspecies by nearly the whole of Sich-
uan. Abies chensiensis ssp. salouenensis
occurs in the S of that province, in NW Yunnan and in the easternmost parts of Xizang [Tibet].
Abies chensiensis ssp. yulongxueshanensis is limited to Yulongxue Shan (‘Jade Dragon Snow Moun-
tain’ after which it was named) and Haba Shan a short distance to the NW, both in NW Yunnan.
The subspecies salouenensis is reported to occur also in Arunachal Pradesh, NE India but no
herbarium records were found to put this on the map. Given its known distribution it could also
be present in the Kachin State of Myanmar [Burma], a region rarely visited by plant collectors
since the travels of F. Kingdon-Ward between 1911 and 1956. Abies chensiensis is a rare species in
most of the S part of its range, usually confined to narrow valleys and ravines following streams
at altitudes between 1600 m and 3200 m and it is usually a minor component of mixed conifer
forests. In the N part, subspecies chensiensis also occupies the lower slopes but occurs from 2000
m to 3500 m, and it is mixed with deciduous angiosperms at the lower and with A. fargesii and
Larix potaninii at the highest elevations.

Map MAJ-74. Abies fabri [n=27, m=27, h=5, 1887–1995]

Abies fabri is endemic to the province of Sichuan

where it occurs in the E and central parts of the
western mountains, from Songpan on the Min
River in the N (ssp. minensis) to Liangshan Yizu
in the S.; to the W it has been collected as far as
Zheduo Shan (Kangding Xian). The type local-
ity of Abies fabri is on Mt. Emei (Emei Shan), a
famous mountain SW of Chengdu. The species
occurs there between 2500 m and 3100 m a.s.l.
in pure stands or in mixed conifer forest, as is
the case in most other localities where this fir
is found. On Zheduo Shan, the westernmost
locality, A. fabri occurs from 3600 m to 4000 m
with Larix potaninii, Picea likiangensis and Juniperus squamata. Near Songpan, the only locality
known for A. fabri ssp. minensis, the high montane to subalpine conifer forests are varied, with
at least three species of Abies, four of Picea, Larix potaninii, Pinus densata and seven species of
Juniperus present in the area. Abies fabri is listed as Vulnerable (VU) on the IUCN Red List due
to decline from logging.
222 mainland asia and japan

Map MAJ-75. Abies delavayi ssp. delavayi

(black) [n=39, m=37, h=11, 1884–2005]; A. dela-
vayi ssp. fansipanensis (red) [n=2, m=2, h=2,
1994–2002]

Abies delavayi is a variable species in which

a number of subspecies and varieties have
been described, but we separate only the
two subspecies here, mainly because there
are insufficient herbarium collections to
separate the varieties meaningfully on the
map and because they are mostly sym-
patric. Abies delavayi ssp. delavayi occurs in NW Yunnan, SE Xizang (mainly in the lower Yarlung
Zangbo drainage), Kachin State of Myanmar [Burma] and Arunachal Pradesh (NE India). For the
latter two areas very few herbarium records are available and A. delavayi is undoubtedly more com-
mon there than the map suggests. The distribution of this subspecies marks the wettest parts of
the Sino-Himalayan mountain system directly influenced by the summer monsoons. Abies delavayi
ssp. delavayi is one of the most subalpine firs of China, commonly occurring between 3000 m and
4000 m (range = 2400–4300 m a.s.l.) together with Picea brachytyla or Picea likiangensis but often
in pure stands up to the tree line. Abies delavayi ssp. fansipanensis is only known from the summit
area (2600–3000 m) of Mt. Fan Si Pan in Lao Cai Province, N Viet Nam. This is ca. 500 km SE of
the nearest populations of the species in Yunnan. Mt. Fan Si Pan (3143 m) is the highest summit in
Viet Nam and culminates the Hoang Lien Range, which connects via the Wuliang Shan and parallel
ranges with NW Yunnan and the Himalaya. Whereas Fan Si Pan at ca. 22.18 N is now the southern-
most locality of Abies in the Eastern Hemisphere, during colder phases of the Pleistocene conifer
forests with this and other species occurred at lower elevations and were probably continuous to
this point or even further S into the Indochina peninsula. Abies delavayi ssp. fansipanensis is listed
as Critically Endangered (CR) on the IUCN Red List.

Map MAJ-76. Abies fargesii var. fargesii

(black) [n=32, m=31, h=7, 1893–1996]; A.
fargesii var. faxoniana (red) [n=28, m=27,
h=8, 1910–1997]; A. fargesii var. sutchuenensis
(yellow) [n=4, m=4, h=2, 1925–1925]

This species is distributed in the N half

of the species-rich area for Abies (see
map MAJ-72 on p. 220) in the mountain-
ous regions of central and N Sichuan,
S Gansu, S Shaanxi, N Chongqing and
W Hubei. Two of the three varieties rec-
ognized in this species are more or less
separated geographically, with A. fargesii var. fargesii in the N of the species range and var. faxo-
niana in the W and S (see map). Both varieties have some outlying populations taking them into
Henan and Xizang [Tibet]/S Sichuan respectively. The third variety, A. fargesii var. sutchuenensis,
has been found in S Gansu but also in the Daba Shan of Chongqing and is therefore sympatric
with var. fargesii. Few herbarium specimens are known of this variety and it is also reported to
occur as expected in NW Sichuan. Abies fargesii occurs at elevations between 2000 m and 4000
m (var. faxoniana between 2600 m and 4000 m) where it forms either pure or mixed conifer
forests with among other species Picea purpurea, P. asperata, P. brachytyla, Larix potaninii, Abies
chensiensis, A. forrestii, A. recurvata, Tsuga chinensis and Taxus chinensis.
 mainland asia and japan 223

Map MAJ-77. Abies forrestii [n=73, m=66,

h=12, 1910–2006]

Abies forrestii is a polymorphic species

which according to Farjon (2001; 2010a)
includes four varieties: A. forrestii var.
forrestii, A. forrestii var. georgei, A. for-
restii var. ferreana and A. forrestii var.
smithii. They are almost entirely sympat-
ric geographically both in terms of distri-
bution range and altitude range (which
does of course not mean that they invari-
ably grow together in the same locality,
although occasionally some do). It is
therefore not informative to map them here on the scale that the Conifer Database geographical
resolution allows. Most of these taxa are also recognized as species by some authors and we may
be dealing with a species complex. The distribution of A. forrestii as circumscribed in Farjon (1990;
2010a) includes SW Sichuan, NW Yunnan and E and SE Xizang [Tibet]. As with some other species
of conifer with this distribution, it is likely to occur in parts of N Myanmar and NE India close
to the border with China, but no herbarium records are available. Abies forrestii is another high
montane to subalpine fir occurring at elevations between 2400 m and 4300 m a.s.l. (commonly
3000–4000 m), either in pure stands or more frequently associated with other species of fir and
spruce and sometimes with Larix potaninii and with various angiosperms at lower altitudes.

Abies forrestii var. georgei near Tienshe Lake, S of Zhongdian, NW Yunnan, China. Photo credit Philippe
de Spoelbergh.

Map MAJ-78. Abies squamata [n=37, m=36, h=10,

1904–1984]

Abies squamata is distributed in W Sichuan

and, sporadically, in E Xizang [Tibet]; some
other outlying localities are in S Gansu and
S Qinghai (Banma Xian) but from Gansu no her-
barium records were available to put these on
the map. This species is exclusively subalpine,
occuring between 3500 m and 4600 m altitude;
its upper limit is as high above sea level as the
224 mainland asia and japan

highest summits in the Alps (except Mont Blanc 4807 m), where the tree line is mostly below
2000 m. While Sichuan is admittedly some 15–16 degrees further S than the Alps, this extremely
high tree line is not even met in the tropics of Asia (on Mt. Kinabalu in Borneo at 6° N ca. 3700
m; in the Highlands of Papua New Guinea at ca. 4–6° S ca. 3800 m) or elsewhere. The influence
of the summer monsoon is the most plausible factor allowing conifer trees to grow well above
4000 m here. Whereas between 3500 and 4000 m A. squamata is usually associated with other firs
and spruces and with Larix potaninii, at the upper limits it grows only with Juniperus squamata,
there a decumbent shrub forming extensive low thickets between the stunted fir trees. This spe-
cies is rare and has been reduced in its abundance by logging; it is listed as Vulnerable (VU) on
the IUCN Red List.

Map MAJ-79. Abies recurvata var. recurvata

(red) [n=15, m=15, h=7, 1903–1999]; A. recur-
vata var. ernestii (black) [n=29, m=29, h=6,
1908–1994]

Abies recurvata is distributed in S Gansu,

N and W Sichuan, E Xizang [Tibet] and
NW Yunnan. The two varieties are more
or less sympatric, but A. recurvata var.
recurvata is more frequently found in
the N part of the species range and
A. recurvata var. ernestii is the more
common variety in the W and S. In
Xizang and Yunnan this species is rare
(records of A. recurvata var. salouenen-
sis are here treated as A. chensiensis ssp.
salouenensis, see map MAJ-73 on p. 221).
Abies recurvata is another high montane to subalpine fir of the high mountains in Sichuan and
surrounding regions, occurring between 2300 m and 3700 m a.s.l. or perhaps even higher. It is
usually a constituent of mixed conifer forests with several species of Abies and Picea co-dominant.
Both varieties of Abies recurvata are listed as Vulnerable (VU) on the IUCN Red List.

Map MAJ-80. Abies holophylla [n=24, m=22, h=8,

1859–1996]

Abies holophylla occurs in the far NE of China (Hei-

longjiang, Jilin and Liaoning provinces), in North
Korea and South Korea, and near Vladivostok in
the Russian Far East. In the N of its range this spe-
cies occurs in lowland hills and on mountains to
ca. 1200 m and is often a common component of
mixed deciduous angiosperm-conifer forest, e.g.
on the lower slopes of the Changbai Shan along
the border between China and North Korea, or it
occurs in mixed conifer forest with Pinus koraien-
sis and Larix gmelinii var. olgensis towards the
Amur River. In the Korean Peninsula it becomes
rare and occurs on isolated mountains, the furtherst S of which is Kumgang-san in Cholla-namdo
Province, South Korea. Abies holophylla is also known from Taechong Island, South Korea, a small
island in the Yellow Sea. On these southern mountains it occurs from 500 m to 1500 m a.s.l. in
mixed deciduous forest.
 mainland asia and japan 225

Map MAJ-81. Abies nephrolepis [n=41, m=36,

h=7, 1896–2007]

Abies nephrolepis has a similar distri-

bution as A. holophhylla (map MAJ-80
on p. 224) but is more widespread and
common. It extends further N into the
interior of Heilongjiang in NE China
and along the Sikhote Alin Range in the
Russian Far East, where it forms exten-
sive forests. The few dots on our map
reflect paucity of herbarium specimens,
not rarity of occurrence. In the W there are isolated populations in Hebei (Xiaowutai Shan,
Zhuolu Xian) and Shanxi (Wutai Shan). In Jilin Province A. nephrolepis is the most common fir
in the Changbai Shan, extending into North Korea and generally occurring above A. holophylla
and the angiosperm-dominated forest. In the Korean Peninsula it occurs on some of the same
mountains as A. holophylla, e.g. Kumgang-san in Cholla-namdo Province, South Korea and on
Taechong Island in the Yellow Sea. It is much more a montane species, with lowland populations
only in the Russian Far East and N Heilongjiang, China but reaching 2400 m on Xiaowutai Shan
(2870 m) W of the capital Beijing.

Map MAJ-82. Abies beshanzuensis (black tri-

angle) [n=4, m=3, h=2, 1975–1991]; A. fanjing-
shanensis (red) [n=3, m=3, h=5, 1981–1986];
A. yuanbaoshanensis (yellow triangle) [n=2,
m=2, h=2, 1977–1978]; A. ziyuanensis (blue)
[n=7, m=7, h=2, 1978–1999]

Four species of Abies occur in isolated

localities in SE China, all are extremely
rare and two are only known from the
type locality. Of these, Abies beshanzuen-
sis at Baishan-zu in the Tung Kung Shan
of S Zhejiang Province must count as the
rarest conifer known to science. Origi-
nally seven trees were found in 1963 but only three survive in the wild, one a large cone-bearing
tree (pictured on p. 226). Abies yuanbaoshanensis is also known from a single locality, Yuanbao
Shan in N Guangxi, but here scattered trees occur on the mountain with Tsuga chinensis in mixed
deciduous forest dominated by oaks between 1700 m and 2050 m a.s.l. Abies fanjingshanensis is
known from two localities in the Fanjing Shan (Wuling Shan) in NE Guizhou. This species occurs
at 2100–2300 m, also with Tsuga chinensis and various angiosperm trees in mixed forest. Abies
ziyuanensis occurs in five locations: in NE Guangxi (Yuanbao Shan), in SW Hunan (Xingni and
Chenbu) and in SW Jiangxi (Jinggang Shan), where in one locality (Pingshui Shan) there is just
one mature tree. This appears to be the species of which more localities, but all with tiny popu-
lations, have been discovered in recent years. The ‘classical’ locality is Yinzhulao Shan in Ziyuan
Xian, NE Guangxi. This species was thought to be closely related to A. beshanzuensis and Chinese
botanist have reduced it to a variety of that species, which if correct would of course take that
species out of the acute danger zone. As accepted here following evidence from DNA (Xiang et
al., 2009), A. fanjingshanensis and A. ziyuanensis meet the IUCN criteria for Endangered (EN) and
A. beshanzuensis and A. yuanbaoshanensis are listed as Critically Endangered (EN). These firs
occur in such limited populations primarily because of climate change, which forced them on
to mountain summits and to compete almost everywhere with more shade tolerant trees. If the
climate becomes still warmer, they may disappear altogether.
226 mainland asia and japan

Abies beshanzuensis in the Tung Kung Shan in S Zheijiang, China. This is the largest and oldest of only
three wild surviving trees, making this the rarest and most threatened conifer in the world. Photo credit
T. R. Dudley.

Podocarpaceae

The family Podocarpaceae is represented in China by Dacrycarpus, Dacrydium, Nageia and Podo-
carpus. For the most part, the species in these genera that occur in China have distributions
predominantly outside that country and are consequently treated in other chapters, particularly
in the chapter Malesia. This leaves only Nageia nagi, Podocarpus chingianus. P. macrophyllus,
P. nakaii and P. tropicalis. Of these, P. nakaii is endemic to Taiwan and is treated with other Tai-
wan endemics seperately; P. chingianus and P. subtropicalis are endemic to mainland China and
Nageia nagi and P. macrophyllus also occur outside its borders.

Nageia

The genus Nageia has an E Asian/Malesian distribution (map MA-54 on p. 323) with most species
occurring in tropical latitudes. One species is an exception to this pattern and accounts for the
extension of the genus into temperate regions of E Asia.

Map MAJ-83. Nageia nagi [n=88, m=78,

h=13, 1863–2011]

Nageia nagi has a very wide distribu-

tion and occurs in SE mainland China,
Hainan, Taiwan and Japan. Its natural
distribution is difficult to establish from
herbarium collections because through-
out its natural range and beyond it has
been widely planted. An attempt to limit
records to those in a forest or mountain
environment has resulted in the map presented here, but there may still be collections included
that come from planted trees near temples or villages. Nageia nagi is certainly indigenous in Hainan
Island, which lies within the tropics but its mountains provide warm-temperate conditions similar
to those in much of the mainland range of this species. In Taiwan natural populations occur in
 mainland asia and japan 227

the N and in the S of the island; it has been planted in many other parts. Nageia nagi also occurs
naturally in the Nansei-Shoto (Ryukyu) archipelago between Taiwan and Japan, on Yakushima and
in Kyushu, Shikoku and S Honshu. In Japan, natural stands are scarce and often consist of only a
few trees, probably the best known and most certainly natural of these are on Yakushima. Nageia
nagi occurs in rich evergreen or semi-deciduous angiosperm forest where its broad leaves enable it
to compete successfully. It is partly sympatric with N. wallichiana but also occurs in more temper-
ate forests where light frost can occasionally occur. In Japan it occurs more often in forests with a
greater component of conifers. Its altitude range is from 200 m to 1300 m.

Nageia nagi near Hongping village, SW Juangxi, China—nat-

ural occurrence or planted? This tree and two others nearby
stood on the steep bank of a river just below a dam in heav-
ily disturbed semi-natural to ruderal vegetation. Remnants
of secondary forest were nearby, but were these trees once
constituents of the forest or were they planted for their ‘fruits’
which alledgedly have medicinal value? This species does
occur naturally in Jiangxi but these trees in a semi-urban set-
ting are not certain evidence of that and so the locality was
not put on the map even though a herbarium specimen was
sampled. Photo credit Aljos Farjon.

Podocarpus

Map MAJ-84. Podocarpus chingianus [n=2, m=1,

h=2, 1925–1926]

Podocarpus chingianus is a poorly known

species of which only two herbarium col-
lections are known with certainty, the type
R. C. Ching 2477 (A) collected in 1925 near
Daxue in Zheijiang Province (mapped) and
C. A. Tso 1477 (A, K) in Jiangsu Province but
with no locality information. A third collec-
tion cited by Hu (1964) from the Harvard
herbaria, W. P. Fang 14261 from Sichuan, has
not been traced. This taxon is listed in the
Chinese Virtual Herbarium under P. macro-
phyllus [var.] chingii with one specimen S.
S. Kuang 11, July 1960 but without location
(or image) at HENU (http://www.cvh.org.cn/
data/index.php accessed 11 Jan 2013). In Flora
of China 4 (Fu et al., 1999: 83) this species is
treated as P. macrophyllus var. chingii occur-
ring in Jiangsu and Zhejiang and (doubtfully) Sichuan. On the IUCN Red List this species is Data
Deficient (DD).
228 mainland asia and japan

Map MAJ-85. Podocarpus subtropicalis [n=5, m=4, h=2,

1887–1942]

This species is also poorly known and originally

described from herbarium material collected on Emei
Shan in Sichuan (E. H. Wilson 3007, October 1903, holo-
type A, isotype K) but earlier collected by E. Faber on
the Jinsha River (“Yangtze-Kiang”) near a monastery
(probably planted) and later also on the mountain Wa
Shan some distance SW of Emei Shan. No other locali-
ties are known; in China this species is not distinguished
from P. neriifolius (map MA-7 on p. 298). It is believed
to be widely planted in SE Asia but there is little or no
documented evidence for that assertion. On the IUCN
Red List this species is listed as Data Deficient (DD).

Map MAJ-86. Podocarpus macrophyl-

lus [n=131, m=124, h=22, 1888–2006]

Two varieties are recognized, Podo-

carpus macrophyllus var. macro-
phyllus and P. macrophyllus var.
maki, but they appear to be almost
wholly sympatric and their distinc-
tion from herbarium specimens
seen as images in the Chinese Vir-
tual Herbarium can be problematic. Therefore they are not separated in the map presented here.
Podocarpus macrophylus is widely distributed from Myanmar [Burma] through mainland China
and Taiwan to Japan. This species is absent on Hainan Island and has not been found in N Viet
Nam, probably these more tropical regions do not have suitable habitat. Podocarpus macrophyllus
is a forest species but also occurs in secondary vegetation (thickets and scrub on mountain slopes
and hill sides), often along streams. It is one of numerous species in a mixed mesophytic forest
formation, largely constituted of deciduous and/or evergreen angiosperm trees and shrubs. Podo-
carpus macrophyllus occurs from near sea level to around 1000 m a.s.l.; in Yunnan it was recorded
(G. Forrest 4665) growing at 2400 m and higher (–3000 m?) as a low shrub. Notably, P. macrophyl-
lus var. maki has been widely planted in China and Japan and has probably been spread by birds
into vegetation in which it may not have occurred previously.

Cupressaceae

The Cupressaceae is a family with 12 genera and 36 species occurring in China and Taiwan. Only
the monospecific genus Metasequoia is endemic to China; 16 species are endemic, 2 of these are
limited to Taiwan. Because of greater adaptation to dry climates than members of other conifer
families, the genus Juniperus occurs widely in the W half of China as well as in the E half and in
Taiwan, making this family the most widely distributed in China.

Cunninghamia

There are two species of Cunninghamia, the phylogenetically most basal genus in the family
(Christenhusz et al., 2011). One is widespread in mainland China but also much planted there and
outside its natural range; the other is rare and has a disjunct distribution in E mainland China,
Taiwan, Viet Nam and Lao PDR.
 mainland asia and japan 229

Map MAJ-87. Cunninghamia lanceo-

lata [n=119, m=111, h=18, 18??–2011]

Cunninghamia lanceolata is prob-

ably endemic in China excluding
Taiwan; as far as known no truly
natural occurrences have been
found in countries bordering
mainland China or in Taiwan. Its
natural distribution within China
is not easy to establish from her-
barium specimen data because
this conifer has probably been
planted in more places, including
afforestation of deforested moun-
tain slopes, than any other species. Large trees in primary forest are extremely rare, perhaps
virtually gone, from logging for many centuries. The map shows its probable distribution, as with
other species in China excluding records of evidently or probably planted trees. As a result, our
map compares reasonably well with the map presented by Ying et al. (2003) in their Atlas of the
Gymnosperms of China, but is considerably less dense (that is, there are fewer dots). The map we
present almost excludes records in Yunnan where this species has been planted in much of the
province but seems to occur naturally only in a few places in the far W and NW and in the E not
far from Guangxi. In Viet Nam it is doubtfully indigenous and not listed in e.g. Nguyen Tien Hiep
et al. (2004) where C. konishii is the only species recognized to occur. Its status in Lao PDR is also
uncertain; no herbarium collections from trees growing wild are available. The natural habitat of
C. lanceolata may have been mesophytic deciduous and/or evergreen forest in the lowlands; it is
at present mostly restricted to steep rocky mountain slopes, crags and other more or less inacces-
sible places where a mixture of smaller angiosperm trees and shrubs and conifers, e.g. Fokienia
hodginsii, Pinus hwangshanensis, P. massoniana, Tsuga chinensis, Taxus chinensis, Torreya grandis,
Cephalotaxus fortunei and locally Cathaya argyrophylla, find a foothold.

Cunninghamia lanceolata on Bijia Shan. Jinggang Shan, Luoxiao Range, Jiangxi, China. Photo credit Aljos
Farjon.
230 mainland asia and japan

Map MAJ-88. Cunninghamia konishii

[n=29, m=22, h=12, 1906–2007]

Cunninghamia konishii is a rare

species with a very disjunct distri-
bution. It was long known to occur
in Taiwan, still its main location,
but more recently it was discov-
ered in Viet Nam and in Lao PDR,
while an earlier collection (W. R.
Price 1238, September 1912 at K) from Fujian, China was more recently identified as this species.
The locality of the latter is uncertain as its transcription on the label does not exactly fit and has
been interpreted as [“Ko-Kle” (prob. Ko-k’eng, = Gekeng)] in the Conifer Database. The second
population of significant size and extent occurs in N Viet Nam and just across the border in Lao
PDR. The southernmost localities are in the N end of the Annam Highlands, the dividing range
between the two countries; two more distant localities are further N along the border. In Taiwan
Cunninghamia konishii is an emergent and sometimes giant tree in rich mixed evergreen forest,
at its lower altitudes around 1000 m predominantly of angiosperms, higher up above 1800 m more
of conifers, to a maximum of ca. 2200 m. In Lao PDR/Viet Nam the altitude range is lower, from
1200 m to 1400(–1660) m and the forest is described as “primary montane mixed coniferous and
angiosperm evergreen forest with Fagaceae, Lauraceae, Magnoliaceae, Fokienia hodginsii, Dacry-
carpus imbricatus.” (Leonid Averyanov on herbarium labels). Cunninghamia konishii is listed as
Endangered (EN) on the IUCN Red List.

Taiwania

Map MAJ-89. Taiwania cryptomeri-

oides [n=66, m=49, h=18, 1868–2006]

Taiwania cryptomerioides is the

only species in its genus (Farjon,
2005, 2010a) and was originally
described and named from Tai-
wan. Although a cultivated tree
was found in SW Yunnan in 1868,
in 1915 it was found growing in the wild in Kachin State, Myanmar [Burma] by J. S. Gamble and
a year later across the border in W Yunnan, China by H. von Handel-Mazzetti. These two very
disjunct populations, separated by ca. 2100 km, have of course led to their separation by some
authors as distinct species. Other records from intermediate areas in China (see e.g. Fu & Jin,
1992 and Ying et al., 2003) are based on introductions of this highly valuable timber tree; none of
these localities refer to the type of primary forest in which T. cryptomerioides is an emergent, slow
growing but long-living tree. In 2001 it was discovered in the S part of the Hoang Lien Range, Lao
Cai Province, Viet Nam. Here a tiny relict population of ca. 100 mature trees survives in remnants
of montane evergreen fagaceous/lauraceous forest of the same type as in Yunnan, but with only
­Fokienia hodginsii as associated conifer. Ecological and historical evidence excludes the possibil-
ity of these having been introduced here. Taiwania cryptomerioides can attain very large dimen-
sions and great age; it is comparable to Fitzroya cupressoides in Chile and Argentina (map SA-32
on p. 451) and to Sequoiadendron giganteum in California, USA (map NA-65 on p. 72) in its general
appearance and ecology. The slow growth of T. cryptomerioides cannot sustain exploitation and
this species is listed as Vulnerable (VU) on the IUCN Red List.
 mainland asia and japan 231

Taiwania cryptomerioides as emer-

gent trees above a mixed canopy of
angiosperms and pines (Pinus sp.) on
a slope in the Gaoligong Shan, NW
Yunnan. Photo credit David Long.

Metasequoia

Map MAJ-90. Metasequoia glyptostroboides

[n=27, m=24, h=17, 1946–2001]

Metasequoia glyptostroboides is another

Chinese example of a monospecific genus,
now endemic to China. It was first identi-
fied as a Pliocene fossil in Japan; during the
Second World War Chinese botanists dis-
covered it alive in the village of Modaoxi
in E Sichuan (now in Chong­qing). Shortly
after, in 1947–48 botanists returned and
found more trees in the area; it became one of the most successful introductions of a Chinese
tree to world-wide horticulture. In the fossil record the genus is recognized to have occurred
from the Late Cretaceous until the Recent in North America and Eurasia; during the warmest
period in the Eocene to well within the Arctic Circle (Stockey et al. in Farjon, 2005a). During the
Pleistocene it retreated to E Asia and was reduced to a single species. In the ‘Metasequoia Valley’
(Shuishaba Valley) in W Hubei, where the living species was found in 1948, a few centuries of
cultivation had left only a few scattered old trees and it took some time until the 1980s before the
species was discovered in more natural habitat in the region. Little if anything now surrvives of
the natural habitat; its optimal environment was probably more or less swampy valley bottoms. It
is now planted all over China (where climate permits) and the world. Natural stands are not easy
to discern from some of these introductions when working from ­herbarium specimens, but our
map is an attempt to include only the former. Most of these ‘genuine’ collections were made by
C. T. Hwa and W. C. Cheng in 1947–48 before any serious planting had started. Metasequoia glyp-
tostroboides occurs naturally (and with certainty) only in Lichuan Shi, westernmost Hubei and in
Shizu Tujiazu Zizhixian and Wan Xian of E Chongqing Municipality (formerly part of Sichuan). It
may also be natural in Longshan Xian and Sangzhi Xian in N Hunan, where Ying et al. (2003) have
put two dots on their map, but no specimens were found from there in the herbaria consulted for
the Conifer Database (7 Chinese, 10 other).
232 mainland asia and japan

Glyptostrobus

Map MAJ-91. Glyptostrobus pensilis [n=51, m=37,

h=10, 17??–2009]

Glyptostrobus pensilis is the only extant species

in its genus. It occurs in SE China, in Viet Nam
and in Lao PDR. In China its natural distribu-
tion is difficult to reconstruct from the avail-
able herbarium records. Its natural habitat,
river banks and low lying marshland between
rivers, has almost everywhere been transformed
to land for agriculture by draining of marshes
and building dams and dikes as well as canals.
Planting of this tree along these water courses
has been going on for many centuries. Interest-
ingly, the localities in Dac Lac Province, S Viet
Nam appear to be in more natural habitat than
the localities in China. Glyptostrobus pensilis is
most common in Guangdong and Fujian and here it is possible that some stands of this tree
along rivers are perhaps replanted on sites where the natural forest with this species was once
cut. We have recorded some other localities in Guangxi, Hunan and Jiangxi that could be semi-
natural (judged from information on herbarium labels) but excluded records e.g. from Sichuan
and Yunnan in the W and Anhui, Jiangsu and Zhejiang in the E, which appear to lie outside its
natural range. In S Viet Nam this species occurs in marshes along upland streams between 500 m
and 700 m a.s.l. It may have occurred in the Red River drainage of N Viet Nam but is now extinct
there. Recently, it was also discovered on the Nakai Plateau in Khammouan Province, Lao PDR,
where is is now known from 5–6 tiny subpopulations (Thomas & Le Page, 2011). Despite its wide
distribution, Glyptostrobus pensilis is listed as Critically Endangered (CR) on the IUCN Red List,
because the Red List is concerned with populations in the wild and there is doubt that any such
populations still exist in China. Those in Lao PDR and Viet Nam are all (very) small and under
threat from the same causes that may have sent this species into Extinct in the Wild (EW) in
China. This is another conifer well known from the fossil record, having occurred from the Late
Cretaceous to the Late Pleistocene in Eurasia and North America; it is now near to extinction
in the wild.

Thuja

There are only two species of Thuja in China, T. koraiensis and T. sutchuenensis. The former occurs
mainly in Korea and only enters China along the border with North Korea in the Changbai Shan
(map MAJ-149 on p. 267). The latter is an extremely narrow endemic in the Daba Shan of E-central
China. The genus is widespread in North America and occurs in Japan and Korea, but somehow
became very rare in China (map MAJ-179 on p. 286). The reverse situation, extinction in North
America and most of Eurasia but survival in China is much more common (Cathaya, Cunningha-
mia, Glyptostrobus, Keteleeria, Metasequoia and possibly Taiwania are examples).
 mainland asia and japan 233

Map MAJ-92. Thuja sutchuenensis [n=6,

m=6, h=6, 1892–2000]

Thuja sutchuenensis was discovered

and described by R. P. Farges at the
beginning of the twentieth century
from an unspecified locality in the
Daba Shan near the city of Cheng-
kou, now in Chongqing Municipal-
ity. It was not found again until 1999
when a Chinese botanical survey for
the newly created municipality col-
lected new specimens (Xiang et al., 2002) and its listing as Extinct in the Wild (EW) could be
reversed. It is now known from Chongqing (Chengkou Co., Kaixian Co.), Sichuan (Wangyuan
Co.), on the southern slope of the Dabashan Mountains, and from Hubei (Baokang Co.), perhaps
now extinct. This very restricted species occupies steep slopes and ridges of limestone mountain
sides between 800 m and 2100 m a.s.l. in mixed angiosperm shrubland and forest. Its conservation
status is Endangered (EN) on the IUCN Red List.

Thuja sutchuenensis in the Daba Shan N

of Chengkou, Chongqing Municipality,
China. Photo credit and © Bedgebury
Pinetum (Daniel Luscombe).
234 mainland asia and japan

Fokienia

Map MAJ-93. Fokienia hodginsii [n=95,

m=88, h=18, 1831–2011]

The genus Fokienia has a single spe-

cies, Fokienia hodginsii, with a wide
distribution in S mainland China, Viet
Nam and in some parts of Lao PDR.
In China it occurs from Yunnan and
S Sichuan, where it is uncommon, E
to Zheijiang. In Viet Nam it is com-
mon through much of the country, as
far S as Chu Yang Sinh National Park
near Da Lat in Lam Dong Province.
In Lao PDR it occurs in the N and is
so far only known from three prov-
inces, mostly near the border with
Viet Nam in the Annam Highlands and in the Pu Bia Massif further W. This conifer can attain a
large size, but such trees due to their very valuable wood have been heavily logged and few remain
in most areas, now almost exclusively in remote mountain locations in Lao PDR and Viet Nam.
There it occurs as an emergent tree in oak-dominated evergreen or semi-deciduous angiosperm
forest, often on steeper slopes or on mountain ridges. Elsewhere, and mostly in China, it is only a
smaller tree occurring with other conifers on rocky ridges or steep slopes at higher altitudes (the
range for the species is 350 m to 2100 m) where it is less exposed to exploitation. This is in part
also its natural habitat, as competition with angiosperms would drive this only moderately shade
tolerant conifer to such exposed sites, especially in regions with evergreen angiosperm forest.
This species is listed as Vulnerable (VU) on the IUCN Red List.

Cupressus

The genus Cupressus is present in China with five species, two of which occur mostly in the
Himalayas and are treated in that section of this chapter. Three species are endemic to mainland
China and these are given maps here. Whereas in North America N of Mexico and in W Eurasia
the species in this genus are all adapted to xeric environments, in China both xeromorphic and
mesomorphic species occur and the endemic species belong (with some variation in one species)
to the latter ecological category. This has of course implications for their distribution.

Map MAJ-94. Cupressus chengiana [n=28, m=24,

h=11, 1900–1996]

Cupressus chengiana is a relatively rare species,

related to C. torulosa and its variety C. torulosa
var. gigantea (map MAJ-27 on p. 191) but geo-
graphically well separated. It occurs in S Gansu
and (mainly) N Sichuan, with a few isolated
populations in the SW and S of that province.
Its principal populations occur in the valleys of
the Bailong River (Gansu and Sichuan), the Min
River and the Dadu River, along the latter NW of
Kangding especially in the Daxue Shan to the W
 mainland asia and japan 235

of the river. The altitude range of C. chengiana is from 1200 m to 2750 m. It can occur in small pure
stands or groves, but is more commonly found on rocky slopes or cliffs among other xeromorphic
shrubs and trees. Cupressus chengiana is listed as Vulnerable (VU) on the IUCN Red List.

Map MAJ-95. Cupressus duclouxiana [n=54, m=21,

h=11, 1890–2000]

Cupressus duclouxiana has a relatively limited

natural distribution in the high mountains and
deep river valleys of S Sichuan and NW Yunnan.
In a few localities in E and W Yunnan distant
from the region where the Nu (Salween), Lan-
cang (Mekong) and Jinsha (Yangtze) Rivers cut
their deep valleys there may also be natural
stands. In Sichuan natural populations also occur
along some western tributaries of the Yalong
River. Elsewhere, and especially in the hills and
mountains around Dali and Kunming, the capi-
tal of Yunnan, this species is apparently introduced and sometimes naturalized, as in the Western
Hills. Most of the early herbarium collections by Europeans (most notably French missionaries)
are from these introductions and these as well as later gatherings are excluded from this map. The
Abbé Delavay understood this well; he notes with his coll. No. 6805: “grande arbre cultivé partout
aux environs d’Yunnan-sen.” Its altitude range in natural populations is probaly between 1900 m
and 3300 m a.s.l. in the deep gorges and on the slopes above the rivers. Cupressus duclouxiana can
occur in nearly pure stands but is more commonly associated with Pinus densata and with Castanea,
Castanopsis and Quercus in mixed evergreen/deciduous oak forest.

Cupressus duclouxiana in the gorge of the Lancang

(Mekong) River NW of Dêqên, NW Yunnan, China.
Photo credit Aljos Farjon.
236 mainland asia and japan

Map MAJ-96. Cupressus funebris

[n=55, m=39, h=9, 1793–2004]

The natural distribution of

Cupressus funebris is even more
difficult to establish than that
of C. duclouxiana; the situation
is not unlike that with C. sem-
pervirens in the Mediterranean
(map EM-29 on p. 159). Planted
over much of China for centuries
in parts of the country where
deforestation is even older and almost complete except for sanctuaries around monasteries and
temples where the Buddhist monks may have either planted or preserved these trees with no
scientific records kept, the difficulty can be imagined. By excluding all herbarium records from
towns and lowlands under agriculture and including those associated with forests and hill or
mountain slopes, we arrived at the map here presented. Although the species is widespread in
S-central China, its natural distribution is much more limited than the map in Ying et al. (2003:
122–123) indicates. These researchers have apparently not tried to make that distinction and
included all herbarium specimens that could be located. From our map it appears that Cupressus
funebris occurs most frequently in the mountainous region of N Guizhou and S Chongqing and
is more sparsely distributed elsewhere, but with perhaps some concentrations in SW Guizhou
and W-central Sichuan. Its natural habitat seems to be more or less open types of deciduous/
evergreen angiosperm forest or mixed forest, but it is now often found in degraded secondary
woodland or scrubland. Here it may colonize, perhaps from planted sources nearby.

Juniperus

The genus Juniperus is present in China with 16 species, of which seven are endemic and four
have wider distributions within China than without. This leaves five species which are treated
in other sections of this chapter. The distribution of the genus in China is similar to that of all
conifers (map MAJ-35 on p. 199) but in Xinjiang and Xizang [Tibet], i.e. the arid and cold W,
they predominate whereas in the more benign climate of the E junipers are much less abundant.
Despite these differences, the distribution of Juniperus in China shows it to be the most successful
genus in terms of biogeography. The genus Abies, with the same number of species in China (map
MAJ-72 on p. 220) has a considerably more limited distribution. Effective long distance dispersal
of the seeds by birds is one factor that has played a significant part, but it is not a sufficient factor,
as other bird dispersed conifer genera such as Podocarpus and Taxus have remained much more
restricted in their distribution. The second factor of importance is adaptation to aridity and low
temperatures, both in situations of low annual rainfall and in exposure to rapid dessication either
at high alpine altitudes or on rocks or sands at lower elevation. Yet some species of Juniperus
occur in mountains with high rainfall under the influence of the monsoon, but in line with many
other conifers, they are capable to survive in mineral deficient soils and thereby avoid competi-
tion from angiosperms with broad leaves which would take out much of the sunlight. It is this
broad array of suitable habitats that is the ultimate cause of the wide distribution of this genus.
 mainland asia and japan 237

Map MAJ-97. The distribution of the genus Juniperus in mainland China and Taiwan.

Map MAJ-98. Juniperus chinensis

var. chinensis (black) [n=91, m=71,
h=17, 1767–2005]; J. chinensis var.
sargentii (red) [n=27, m=26, h=8,
1861–1975]; J. chinensis var. tsuku-
siensis (yellow) [n=5, m=5, h=3,
1928–1994]

Juniperus chinensis is widely

ditstributed from Sichuan
and Yunnan in the W through
much of China, including Tai-
wan, to Jilin in the NE, and
in Korea, Japan and Sakhalin
Island in the Russian Far East.
The three varieties recognized
have different ranges, but only
J. chinensis var. tsukusiensis is mostly allopatric, occurring in NE Taiwan on a few ocean-facing
cliff sides, where it is the only variety in the wild, and on Yakushima in Japan. Juniperus chinensis
var. sargentii, another shrub form of the species, occurs in Jilin and Heilongjiang, NE China, in
Korea, Japan and Sakhalin. It is the northernmost variety, but in Honshu and Korea J. chinensis
var. chinensis is also found; this variety is the only form present in most of mainland China. The
two widespread varieties occur in different habitats, so they are not sympatric on the smaller
landscape scale. The tree form, J. chinensis var. chinensis, is also very widely planted especially
in the more northern, colder parts of its range, where, together with Platycladus orientalis (map
MAJ-107 on p. 241) it is the most common traditional conifer tree in temple grounds and other
historical sites (e.g. the ‘Forbidden City’ in Beijing). In a few localities this widespread variety
forms groves of tall trees (e.g. in S Gansu), or it is mixed with pines and deciduous angiosperms
at canopy level. It is much more common, under conditions largely determined by man’s agricul-
tural practices, in secondary vegetation, on open, rocky slopes. The altitude range is (100–)1400–
2400(–2700) m a.s.l. High montane varieties J. chinensis var. sargentii and var. tsukusiensis occur
on rocky outcrops and amongst boulders and have attained a decumbent habit.
238 mainland asia and japan

Map MAJ-99. Juniperus convallium [n=70, m=68,

h=14, 1900–1998]

Juniperus convallium occurs in SE Xixang

[Tibet], S Qinghai, W Sichuan and S Gansu.
The two varieties recognized (Farjon, 2010a) are
sympatric and not separated on the map pre-
sented here. In 1997–98 extensive collecting in
Xizang by G. and S. Miehe (specimens at GOET
and K) has substantially added to our knowledge of the distribution of this species, which not
only occurs in the Lancang (Mekong) and Jinsha (Yangtze) drainages but also to the W in the
Yarlung Zangbo drainage, where it was collected as far W as Nyemo. Juniperus convallium is a
high montane to subalpine species with an altitude range from 2200 m to 4430 m and occurs on
S-SW facing slopes in drier and warmer parts of conifer forest or degraded woodland. It usually
stays below J. tibetica but can occur in some localities mixed with it.

Map MAJ-100. Juniperus tibetica [n=124, m=115,

h=16, 1880–2005]

Juniperus tibetica has a similar but wider

distribution than J. convallium and occurs
in S-SE Xizang [Tibet], extreme NW Yun-
nan, W Sichuan, E Qinghai and W Gansu.
In Xizang it follows the Yarlung Zangbo
River and some of its tributaries as far
W as Lhading Gompa (86° 52’ E) beyond
which point trees become very rare. Juni-
perus tibetica forms groves or small forests
at high altitudes, often together with J. con-
vallium but extending higher than that species; the altitude range is 2600–4780(–4900) m a.s.l. In
Xizang it mixes locally in open woodlands with Cupressus torulosa var. gigantea (map MAJ-27 on
p. 191) and both species can attain large size and presumably great age. These woods are often
overgrazed by yaks and being transformed into parkland-type vegetation with mats of Cyperaceae
where regeneration is difficult. This species is listed as Vulnerable (VU) on the IUCN Red List.

Map MAJ-101. Juniperus saltuaria [n=64, m=54,

h=14, 1885–2001]

Juniperus saltuaria occurs in SE Xizang

[Tibet], NW Yunnan, W Sichuan and
S Gansu. Its distribution is similar to that
of J. tibetica (map. MAJ-100 on p. 238) but
does not extend as far W in Xizang and it
is probably less common there but may
have been less frequently collected for her-
baria. It is reported from SE Qinghai but
no herbarium collections were available;
according to the map presented in Ying et al. (2003) it is present in a location in the far E of that
province. The altitude range of this species is from 2100 m to 4600 m a.s.l. and it occurs in more
or less open conifer forests with Larix potaninii and Abies and Picea and in more or less pure juni-
per woods with J. tibetica and J. convallium and/or the decumbent forms J. pingii var. wilsonii and
J. squamata up to the tree line and beyond.
 mainland asia and japan 239

Map MAJ-102. Juniperus komarovii (black) [n=27, m=25, h=12,

1922–2001]; J. przewalskii (red) [n=54, m=41, h=13, 1872–2007]

Juniperus komarovii and J. przewalskii are similar species

and both are closely related to J. tibetica, J. convallium and
J. saltuaria (Farjon, 2005; Adams, 2011). This group of spe-
cies occupies the E Tibetan Plateau (in its widest sense) and
J. komarovii and J. przewalskii are distributed in the NE section
of its combined range. Juniperus komarovii occurs mostly to
the S of J. przewalskii in NW Sichuan from the Daxue Shan in
the W to SW Gansu and sporadically in SE Qinghai. Juniperus
przewalskii occurs (sporadically) in NW Sichuan, in S Gansu
and in most of E Qinghai, where it reaches its N limit in the
Qilian Shan which on its N slopes drops off steeply into the
Gobi Desert. Juniper trees are absent in a large intermoun-
tain depression centred around Qinghai Hu (Koko-nur), the
largest (but shrinking) lake in China which is saline and lies
at 3205 m a.s.l. This large basin is mostly treeless desert. Juni-
perus komarovii and J. przewalskii are partly sympatric with
Picea asperata (map MAJ-49 on p. 208) and P. crassifolia (map MAJ-56 on p. 211) but whereas the
spruces occupy N-facing slopes, the junipers dominate on the S-facing slopes. Both species are
high altitude trees or large shrubs occurring between ca. 2000 m and >4000 m.

Map MAJ-103. Juniperus pingii var. pingii + var. chengii (black) [n=22, m=21, h=6, 1885–2000]; J. pingii var.
wilsonii + var. miehei (red) [n=101, m=98, h=10, 1889–2003]

Juniperus pingii is a variable species which may well form hybrid swarms with J. squamata (they
often occur together as on Baima Shan, NW Yunnan) and Adams (2011) treats J. pingii var. wilsonii
(excl. J. pingii var. carinata) as J. squamata f. wilsonii. The most widespread variety of this spe-
cies is var. wilsonii, the decumbent subalpine shrub; the small tree or large erect shrub J. pingii
var. pingii is much more restricted occurring in W and S Sichuan and NW Yunnan. The variet-
ies chengii (small tree) and miehei (decumbent shrub) are too poorly known to be separated on
the map and so are included with the taxa they most resemble both morphologically and geo-
graphically. Juniperus pingii var. wilsonii is widely distributed and occurs from a few localities in
W Hubei, S Shaanxi and S Gansu through W Sichuan, NW Yunnan and SE and S Xizang [Tibet],
westwards to the upper drainage of the Yarlong Zangbo River, where S of Saga, along the road
to Gyirong it reaches its westernmost locality at 29° 14’ N 85° 19’ E as well as its highest known
altitude, 4850 m. [This population was described as var. miehei in Farjon, 2005a: 346.]
240 mainland asia and japan

Juniperus pingii var. wilsonii (fore-

ground and some of the low patches
on a slope in far left of photograph)
with Larix potaninii and Picea liki-
angensis at ca. 4000 m, Baima Shan
Nature Reserve, Dêqên Xian, NW
Yunnan. Photo credit Aljos Farjon.

Map MAJ-104. Juniperus formosana [n=136,

m=131, h=24, 1886–2001]

Juniperus formosana has the widest dis-

tribution of the genus in China. It occurs
from Taiwan in the E across mainland
China to NW Yunnan, W-central Sich-
uan, S Gansu and extreme E Qing-
hai. It is uncommon or rare in Hunan,
Guangdong (no herbarium records
found) and Guangxi and absent in
S-central Yunnan. The map presented
by Ying et al. (2003: 136–137) has dots in
N Guangdong and more dots than our map in Guangxi, but these records were not present in
the datasets accessed from herbaria participating in the Chinese Virtual Herbarium (http://www
.cvh.org.cn/data/index.php accessed 15 Jan 2013). This species is most commonly found in open
scrubland in rocky or gravelly places on hills and mountain slopes, in river beds and ravines. At
higher altitudes in the W of its range it is commonly present in open areas of conifer forest, espe-
cially associated with Pinus as these forests are lighter than those dominated by Abies or Picea.
Juniperus formosana has a large altitude range from 400 m to 3830 m and at higher altiitudes it
may grow with (and be confused with) J. squamata. For this reason the Conifer Database only
accepted specimens seen, either really or virtually (i.e. images).

Calocedrus

The genus Calocedrus has four species and is disjunctly distributed in the Northern Hemisphere,
with one species in W North America (map NA-90 on p. 86) and three species in SE Asia. This dis-
junction is the result of vicariance distribution, not long distance dispersal; there is a fossil record
from the Oligocene-Early Miocene in Central and SE Europe and from the Pliocene in Poland
(Kvaček, 1999). In the Miocene it occurred in Idaho, so it may have had a wider distribution away
from the W coast in North America. A migration route across the Bering Land Bridge is of course
a possibility. Chen et al. (2009) estimated a divergence time in the Oligocene (ca. 25 Ma) for the
split between C. decurrens (North America) and C. macrolepis + C. formosana, representing the
Asian clade. Their results are interpreted as “geohistorical vicariance events” in accordance with
the fossil record. Of the Asian species, Calocedrus formosana (map. MAJ-131 on p. 254) is endemic
to Taiwan, C. rupestris (map. MAJ-141 on p. 261) occurs in Viet Nam and Guangxi, China and C.
macrolepis is distributed in mainland China, Hainan Island and Indochina.
 mainland asia and japan 241

Map MAJ-105. The distribution of Calocedrus in North America and SE Asia.

Map MAJ-106. Calocedrus macrolepis [n=54, m=52, h=14,

1868–2009]

Calocedrus macrolepis occurs in Myanmar [Burma]

close to the border with China, in Yunnan, Guizhou,
Guangxi and Hainan (China), in Viet Nam, Lao PDR
and Thailand. This species has a scattered distribution
with gaps between known localities, e.g. in Viet Nam
and in E Yunnan, Guizhou and Guangxi. It also seems
to be very rare in Lao PDR and Thailand, although
here insufficient data could be another cause for the
few dots on the map. Calocedrus macrolepis can be
locally common in montane mixed evergreen conifer-
broad-leaved forest dominated by Fagaceae and with
scattered conifers, e.g. Cunninghamia, Dacrycarpus,
Keteleeria and Pinus; it is also often planted in road-
sides and field margins. The altitudinal range is from
ca. 800 m to 2000 m a.s.l.

Platycladus

Map MAJ-107. Platycladus orientalis [n=48,

m=17, h=16, 1700–1996]

The monospecific genus Platycladus is

not closely related to Thuja (Linnaeus
described and named it as belonging to
Thuja) but to Microbiota decussata (map
MAJ-148 on p. 267). The natural distribu-
tion of Platycladus orientalis is one of the
greatest conundrums of the biogeogra-
phy of Chinese trees. It is one of the most widely planted conifers in China and has found its way
with the help of people across most of temperate Asia and beyond. It has naturalized in many
places, from Korea to Iran, while even if apparently spontaneous it is usually found in secondary
vegetation or much disturbed forest. Platycladus orientalis is probably endemic to China and most
likely has a limited natural distribution in the mountains WSW of the capital Beijing (where it is
planted everywhere in the parks and surrounding hills). It occurs naturally in SE Gansu, Shaanxi,
Shanxi, Henan and Hebei, possibly in nearby parts of Nei Monggol (Inner Mongolia). References
to its occurrence in North Korea and/or South Korea, as well as in the Russian Far East (e.g. in
Farjon, 2005, 2010a) could not be substantiated with herbarium records of unambiguously wild
growing indigenous trees; introduction there seems more likely.
242 mainland asia and japan

Cephalotaxaceae

The family Cephalotaxaceae consists of a single genus, Cephalotaxus, with eight species. Although
found nested within a larger family Taxaceae in several phylogenetic studies using DNA sequence
data, the most recent of such analyses (S. Mathews, unpublished data) using data from six genes and
wide taxon sampling found Cephalotaxus to be the sister group of Taxaceae as circumscribed from
morphological evidence and accepted in Farjon (2010a) and other compilations of gymnosperm
taxonomy. We therefore deviate here from the classification of gymnosperms by Christenhusz et al.
(2011) otherwise followed in this section of the Atlas. The distribution of this family is E Asian, with
seven of the eight species present in China, but a few extending the family eastwards to Japan and in
a SW direction to NE India and Indochina. As with many other E Asian conifer genera, Cephalotaxus
had a much wider distribution in the geological past including Europe, Greenland and W North
America (Florin, 1963), with the earliest known fossils interpreted as Cephalotaxus foliage shoots
from the Jurassic in S Sweden and E Greenland. Molecular estimates of divergence time place the
split between Cephalotaxus and Taxaceae at 147.79 Ma, i.e. in the Late Jurassic (S. Mathews, unpub-
lished data). In the Pliocene (5.3–2.6 Ma) Cephalotaxus still existed in California and Europe. Fossils
have never been found in E North America or in continental Asia outside its present distribution. A
more detailed analysis which samples all eight species will be required to fully resolve the relation-
ships among extant species. If this were to uphold the present results, C. harringtonii is sister to the
other species. This species occurs in China, Korea and Japan; the fossil record of the genus in Japan
goes back to the Miocene, which is again in congruence with molecular dating estimates. However,
given the wider fossil record, it is clear that the true phylogeny of Cephalotaxaceae cannot be recon-
structed from a few surviving species in one corner of the former range. Cephalotaxus harringtonii
(map MAJ-188 on p. 290) occurs outside China and C. mannii (map MAJ-142 on p. 262) only partly
within China; the other six species are treated here: five of them are endemic to China.

Map MAJ-108. The present global distri-

bution of the species of Cephalotaxaceae/
Cephalotaxus.

Cephalotaxus

Map MAJ-109. Cephalotaxus fortunei var. fortunei

(black) [n=136, m=128, h=12, 1889–2006]; C. fortunei
var. alpina (red) [n=28, m=24, h=7, 1897–2008]

Cephalotaxus fortunei has a very wide distri-

bution in China and occurs from just inside
the border in Myanmar [Burma] to the coast
of the East China Sea in Zhejiang Province.
In the N it extends as far as S Gansu and
S Shaanxi, in the S it reaches close to the coast
 mainland asia and japan 243

in E Guangdong and the far S of Yunnan. Most of this wide range is occupied by C. fortunei
var. fortunei, but within the W parts of this range var. alpina also occurs. This latter variety has
been found in S Gansu, central and S Sichuan and Yunnan. Cephalotaxus fortunei grows as an
under-storey tree or shrub in mixed angiosperm forests, in mixed conifer-angiosperm forests and
in conifer forests (Abies, Picea and Larix), with var. alpina only in the conifer forests. It is also
common as a shrub or small tree in open thickets and on roadsides in secondary vegetation, so
it appears to spread and establish easily. Its altitudinal range is great and extends from 200 m to
3700 m a.s.l., with C. fortunei var. alpina at the higher range between (1100–)1800 m and 3700 m.

Cephalotaxus fortunei var. fortunei in winter

deciduous secondary woodland at Mogan
Shan, Zheijiang Province, China. Photo credit
Aljos Farjon.

Map MAJ-110. Cephalotaxus oliveri [n=34, m=33,

h=14, 1889–2001]

Cephalotaxus oliveri has a more restricted

distribution than C. fortunei and is much less
common. Some populations, e.g. in S Yunnan
near the Vietnamese border and in W-central
Sichuan, are isolated. A well-known popula-
tion occurs on Emei Shan SW of Chengdu
in Sichuan. Cephalotaxus oliveri is an under-
storey shrub in a variety of forest types at
altitudes between 300 m and 1800 m; it does
not thrive in full sunlight as C. fortunei and therefore is more vulnerable to deforestation or forest
degradation. As a consequence it has become rare in many areas and is listed as Vulnerable (VU)
on the IUCN Red List.

Map MAJ-111. Cephalotaxus sinensis [n=91, m=87,

h=11, 1886–1996]

Cephalotaxus sinensis has a wide distribution

similar to C. fortunei (map MAJ-109 on p. 242
but unlike that species also occurs in Hainan
Island and somewhat further N in Shaanxi
and Henan Provinces, i.e. its N-S extent is
greater. In NW Yunnan it does not seem to
occur westwards beyond the deep Langcang
River valley, where C. fortunei continues to
the border with Myanmar [Burma]. The
range of habitats in which C. sinensis occurs
244 mainland asia and japan

is also similar; in the E it occurs generally at lower altitudes, from ca. 200 m to 1700 m; in the
W it reaches >3000 m in Sichuan and Yunnan. The forest types in which it occurs also change
from E to W, from predominantly angiosperms to predominantly conifers. In either type of forest
C. sinensis is usually a small under-storey tree, but it will also grow in shrubby thickets where it
will itself remain a shrub.

Map MAJ-112. Cephalotaxus latifolia [n=17, m=17, h=7,

1928–1996]

Cephalotaxus latifolia occurs in Chongqing, NW

Fujian, N Guangdong, NE Guangxi, SE Guizhou, SW
Hubei, Hunan and W Jiangxi. No herbarium specimen
records could be accessed for Fujian, Guangdong,
Hunan and Jiangxi. In Hunan it may only occur close
to the border with Guangxi as there are several records
from NE Guangxi close to that border. Difficulty with
identification of the various species of Cephalota­
xus means that only records from specimens seen
(real or virtual) could be entered in the Conifer Data-
base unless the identifier is known to have good
knowledge of the species. Localities in the provinces
for which we have no records seem to be outliers
(Ying et al., 2003: 180–181) based on specimens from
herbaria to which we had no access. Cephalotaxus lati-
folia occurs in mountains at altitudes between 900 m and 2400 m. It grows in secondary vegeta-
tion (“thickets” according to the Flora of China, Vol. 4: 86, Fu et al., 1999) usually forming a shrub.
It may well be more common than previously thought; recent inventories in the new Chongqing
Municipality (an administrative split from E Sichuan Province) have found it there in several new
locations.

Map MAJ-113. Cephalotaxus hainanensis [n=6, m=6, h=8,

1922–1979]

Cephalotaxus hainanensis is endemic to Hainan Island,

where it occurs in the following mountains: Jianfeng
Ling, Limu Ling, Wuzhi Shan, Dawang Ling and Diao
Luo Shan. Four of these have herbarium records and
are shown on the map; we are missing a record for
Dawang Ling (19° 29’ N 109° 44’ E) and this is shown
in red. Cephalotaxus hainanensis occurs in mixed
warm temperate to subtropical rainforests (evergreen
broad-leaved forest) in mountainous areas of the
island, where it attains tree habit and size (10–20 m).
This species occurs from near sea level up to about
1700 m, but usually between 500 m and 1100 m. It can
sometimes be the dominant tree in the community but is usually associated with Dacrydium
pectinatum, Dacrycarpus imbricatus, Nageia nagi, Podocarpus neriifolius and various angiosperms
among which can be palms. Cephalotaxus hainanensis is listed as Endangered (EN) on the IUCN
Red List.
 mainland asia and japan 245

Map MAJ-114. Cephalotaxus lanceolata [n=5,

m=5, h=4, 1959–2006]

Cephalotaxus lanceolata has an extremely

limited distribution in a few localities on
the upper Dulong River W of Gongshan,
close to the Myanmar [Burma] border. It
was long known from the type collection
only, gathered in 1959 by G. M. Feng and
kept in the Chinese herbaria KUN and
PE. In 1990 and in 2006 joint Chinese/
American teams collected it again in the vicinity of Nangza near the tiny village of Maku on
the same river, only a few km from the international border. The locality of the type collection
was not precisely indicated and its placement as the most eastern dot on our map is tentative,
it could have been closer to where it was later collected. It is possible that this rare species is
also present across the divide in Myanmar, but no investigations have been undertaken there.
Cephalotaxus lanceolata occurs scattered in evergreen angiosperm forest at ca. 1450–1900 m
elevation. Due to its rarity and inferred decline from logging in the area this species is listed as
Endangered (EN) on the IUCN Red List.

Taxaceae

Four of the five genera in Taxaceae are present in China, the only genus that is missing is the
monospecific genus Austrotaxus, endemic to New Caledonia (map AU-94 on p. 405). Pseudotaxus,
another monospecific genus, is endemic to China and two genera, Amentotaxus and Torreya, have
their main distributions here in terms of species diversity (Torreya) or area of occupancy (AOO)
in the case of Amentotaxus. The sister group of Taxaceae, Cephalotaxus (Cephalotaxaceae) is also
centred in China. However, Taxus is distributed across the Northern Hemisphere and in Malesia
crosses the Equator. When we include evidence from the fossil record we find that most of these
genera were also widespread far beyond China, so that the generic diversity of China is a case of
survival in that region of E Asia, not of origin. A map of the family is therefore not given here but
in the chapter Global and trans-continental distributions.

Amentotaxus

The genus Amentotaxus has six species, three species in NE India and Indochina (see maps MAJ-
34 and MAJ-143 on p. 196 and p. 263) and three in mainland China and Taiwan, of these A. for-
mosana is endemic to Taiwan and is treated separately. Amentotaxus has its main distribution in
S-central China, with several outlying populations involving different species. The northernmost
locations are in S Gansu near the border with Sichuan, to the E it extends to Taiwan, to the S
as far as the Ngoc Linh Mountains in Viet Nam (15° 04’ N) and to the W to Arunachal Pradesh
in NE India. Large gaps occur in the W, where one or two localities are known in S-central
Sichuan, one in central Yunnan, one in SE Xizang [Tibet] in the bend of the Yarlung Zangbo/
Brahmaputra River and a few in Arunachal Pradesh. Another gap may exist between the east-
ernmost occurrences in mainland China and Taiwan (see under A. argotaenia map MAJ-116 on
p. 246 for caveat). Amentotaxus occurs in the Cenozoic fossil record of North America and Europe;
in the latter continent it was present from the Palaeocene to the Upper Miocene (Ferguson
et al., 2009).

Map MAJ-100. Ju
246 mainland asia and japan

Map MAJ-115. The present global dis-

tribution of the genus Amentotaxus.

Map MAJ-116. Amentotaxus argotae-

nia [n=81, m=77, h=17, 1882–2011]

Amentotaxus argotaenia has by far

the widest distribution of all spe-
cies with nearly the same extent
of occurrence (EOO) as the entire
genus. The genus is centred in
S-central China but extends into
N Viet Nam and adjacent parts
of Lao PDR. A small population
occurs in the extreme S of Tai-
wan at O-luan-pi (“Formosa South
Cape”). Another isolated population is in the W in the bend of the Yarlung Zangbo River in SE
Xizang [Tibet]. The farthest N and S occurrences of this species coincide with the geographical
limits of the genus (see above under the genus and map MAJ-115). Amentotaxus argotaenia is
reported from further E on the mainland in Anhui, Jiangsu and S Zhejiang in Flora of China 4 (Fu
et al., 1999) and Farjon (2010a) (see also the map in Ying et al., 2003: 186–187) but the 18 herbaria
(9 Chinese) consulted for this species in the Conifer Database appear to hold no specimens from
these provinces. Amentotaxus argotaenia occurs on limestone as well as sandstone, shale or gran-
ite, and in ravines, on steep slopes or cliffs, on summits and ridges and in mountain forests along
shady stream banks. The altitude range is between 600 m and 1100 m a.s.l.

Amentotaxus argotaenia sapling on the steep bank of a rocky stream near Xiping Village, Jinggangshan, SW
Jiangxi, China. Photo credit Aljos Farjon.
 mainland asia and japan 247

Map MAJ-117. Amentotaxus yunnanensis [n=19,

m=19, h=10, 1932–2008]

Amentotaxus yunnanensis occurs in SW Gui-

zhou (Xingyi), SE Yunnan (Funing, Malipo,
Maquan and Xichou counties), in N Viet Nam
(Bac Can, Ha Giang, Ha Tuyen, Lao Cai, Nghe
An and Thanh Hoa provinces) and in Lao PDR
(Houaphan province). This distribution largely
coincides with areas of karst limestone forma-
tions and indeed it is most common on the
steep ridges and summits of ‘sugarloaf’ lime-
stone mountains and pinnacles. In this extreme
habitat A. yunnanensis mixes with other coni-
fers, e.g. Fokienia hodginsii, Xanthocyparis vietnamensis, Pseudotsuga sinensis, Tsuga chinensis,
Podocarpus neriifolius, Dacrydium elatum and Taxus chinensis, as well as with small-leaved angio-
sperms. In a few localities A. yunnanensis has been found on substrates derived from acidic rocks
(granite, gneiss) where it is sometimes a large tree amongst angiosperms, with only a few associ-
ated conifers e.g. Cephalotaxus mannii, Dacrycarpus imbricatus and Nageia wallichiana. This spe-
cies is listed as Vulnerable (VU) on the IUCN Red List.

Torreya

The genus Torreya has six species, two of which occur in North America and four in Asia, a pat-
tern that is not unusual in conifers at the genus rank (e.g. Pseudotsuga, Tsuga, Calocedrus, Thuja)
and common in angiosperm trees. The gap, Europe, is in nearly all cases caused by Pleistocene
extinctions. The genus Torreya has been present in Europe for a very long time, from the Middle
Jurassic (ca. 165 Ma) to the Pleistocene (<1 Ma) (Florin, 1963; Li, 1995). In North America the old-
est fossils date from the Lower Cretaceous. In the Cenozoic it had a wider distribution in North
America (particularly W North America) than at present, so retreat has occurred there too, but
not to the point of extinction, although T. taxifolia (map NA-95 on p. 89) has come very close.
Fossil evidence from the Oligocene has shown the genus to have occurred further N in China
than at present, but if this implies a range reduction in E Asia it has been minimal in com-
parison. With Amentotaxus the genus Torreya forms a sister clade to the remainder of Taxaceae
(S. Mathews, unpublished data). Within the genus Torreya two clades emerged, one with the two
North American species and the other with the Asian species (Li et al., 2001). This would appear
to suggest a vicariance event splitting the two groups, but the estimated young age of extant spe-
cies relative to the time since North America separated from Eurasia makes this scenario unlikely.
For a meaningful phylogeny and biogeographic history of the genus Torreya it will be necessary
to integrate data from the long fossil record (Li et al., 2001).

Map MAJ-118. The present distribution of the genus Torreya in W and E North America and in E Asia.
248 mainland asia and japan

Map MAJ-119. Torreya fargesii

var. fargesii (black) [n=35, m=34,
h=10, 1889–1984]; T. fargesii var.
yunnanensis (red) [n=24, m=22,
h=9, 1914–2002]

The two varieties, Torreya

fargesii var. fargesii and var.
yunnanensis, are well sepa-
rated geographically. Tor-
reya fargesii var. fargesii is
scattered in Sichuan on its W limits and in N Hubei, N Jiangxi and S Anhui on its E limits; the
area in which it is more frequent extends from SE Gansu and SW Shaanxi through Chongqing
and SE Shaanxi to W Hubei. This is the range of mountains (Micang Shan, Daba Shan, Wu Shan)
bordering the Sichuan (Red) Basin to the N and NE. Torreya fargesii var. yunnanensis is limited to
NW Yunnan, where it occurs in the Gongshan, Lijiang, Nujiang Lisu and Weixi Lisu Prefectures.
Torreya fargesii replaces T. grandis westwards, but the two species are partially sympatric. Tor-
reya fargesii occurs in coniferous, angiosperm dominated and mixed forests as an under-storey
shrub or a small to medium sized tree, in mountains from 700 m to 3400 m a.s.l. Due to deforesta-
tion both varieties are listed as Endangered (EN) on the IUCN Red List.

Map MAJ-120. Torreya grandis var.

grandis (black) [n=50, m=49, h=10,
1855–2002]; T. grandis var. jiulongshan-
ensis (red) [n=3, m=3, h=2, 1959–1991]

Torreya grandis is distributed in the

SE of China and occurs in Anhui,
Fujian, Guizhou, Guangdong,
Henan, Hubei, Hunan, Jiangxi,
Sichuan and Zhejiang provinces.
The record for Sichuan is based on
a recently (2006) described species,
T. parvifolia, which was considered
synonymous with T. grandis (Far-
jon, 2010a). Herbarium records
were not available via the Chinese
Virtual Herbarium (http://www
.cvh.org.cn/data/index.php accessed 17 Jan 2013) and so it cannot be placed on the map. West-
wards this species is gradually replaced by T. fargesii (map MAJ-119) and there is an issue with the
correct identification of specimens especially in the areas where the two species ranges overlap.
Torreya grandis var. jiulongshanensis is only known from the Jiulong Shan in S Zheijiang, where it
has been found in two localities. Torreya grandis occurs usually scattered among deciduous and/
or evergreen angiosperm trees in forests at altitudes between 200 m and 1400 m with the decidu-
ous trees in the forest increasing with altitude and latitude.
 mainland asia and japan 249

Torreya grandis in (mostly) deciduous angio-

sperm forest on Tianmu Shan, Zhejiang prov-
ince, China. Almost all species in Taxaceae are
shade tolerant, and Torreya grandis is a good
example, here pictured as a relatively young
tree slowly elevating its crown though the
canopy of a partially evergreen angiosperm
dominated forest. The season is December 2010.
This species does not form extensive stands,
although several trees can occur together, but
rather disperses randomly through the forest
as solitary individuals. As is typical also in this
family, trees of T. grandis grow more slowly but
live longer than most if not all the surround-
ing angiosperm tree species. Forest fires are
uncommon on this mountain which receives
much rainfall especially in summer. Photo
credit Aljos Farjon.

Map MAJ-121. Torreya jackii [n=8, m=7, h=7,

1924–1991]

Torreya jackii is a rare species with a lim-

ited distribution in E China, it occurs in
N Fujian (Taining, Puchen), NE Jiangxi
and Zhejiang. Localities in Fujian Prov-
ince could not be mapped because we
had no access to relevant herbarium
specimens. The map in Ying et al. (2003)
has three symbols in Fujian and several
in Zhejiang we do not have, but it misses
the two in Jiangxi we have. These Chi-
nese workers have had access to some
herbaria that are not represented in the
Conifer Database. There may also be
an issue with records from cultivation;
when in Zhejiang Province in December
2010 one of us (AF) was shown several trees in cultivation but encountered none in the wild.
Despite this, the map presented here is undoubtedly incomplete. Torreya jackii occurs in ever-
green angiosperm forest, along streams, on steep slopes in shade or in secondary vegetation usu-
ally near moisture. Its altitudinal range is between 120 m and 1320 m a.s.l. Due to deforestation
this species is listed as Endangered (EN) on the IUCN Red List.
250 mainland asia and japan

Pseudotaxus

Map MAJ-122. Pseudotaxus chienii [n=27,

m=27, h=8, 1933–2011]

The genus Pseudotaxus is monospecific

and P. chienii is endemic to SE China.
Although very similar to Taxus except
for its white arils around the seeds and
silvery white stomata bands on the
leaves, DNA sequence data from six
genes have confirmed its separation
from Taxus as it resolved as the sister
to all 10 species of Taxus in a recent
phylogenetic analysis (S. Mathews,
unpublished data). Pseudotaxus chienii
occurs in N Fujian, N Guangdong (Nan Ling Shan, Ruyian), Guangxi, N and SW Hunan, SW
and NE Jiangxi and SW Zhejiang. Until very recently, this species was not present in Western
herbaria (it is now in E, collected by K. D. Rushforth in 2011 and in K collected by A. Farjon
in 2009). The map is therefore largely based on data from Chinese herbaria accessed via the
Chinese Virtual Herbarium (http://www.cvh.org.cn/data/index.php), with a few exceptions
of specimens seen in PE on visits by AF. Although there are some minor differences, the map
in Ying et al. (2003) agrees with ours and both show the scattered pattern of distribution of
P. chienii in SE China. This species occupies a spatially limited type of habitat: rock faces above
streams or on shady, moist to dripping sides of ravines and mountains at altitudes between
700 m and 1500 m. Although these are in principle plentiful in SE China, dispersal to these places
by birds and establishment of seedlings may be rare random events. If sites contain a single shrub,
cross pollination may also be difficult. Deforestation affects this species by exposing it, drying out
its habitat. Pseudotaxus chienii is listed as Vulnerable (VU) on the IUCN Red List.

Pseudotaxus chienii in Mang

Shan Park, S Hunan, China.
Photo credit Aljos Farjon.
 mainland asia and japan 251

Taxus

There are four species of Taxus in China, T. chinensis, T. cuspidata, T. mairei and T. wallichiana. Of
these, T. cuspidata (map MAJ-189 on p. 291) and T. wallichiana (map MAJ-33 on p. 195) are treated
in other sections of this chapter.

Map MAJ-123. Taxus chinensis [n=61,

m=59, h=14, 1889–2008]

The distribution of Taxus chinensis

is scattered across a large part of
China and extends into N Viet Nam.
This species is rare or absent in the
easternmost provinces where it is
replaced by T. mairei (map MAJ-
124). In the W it occurs in S Gansu
and Sichuan but there T. mairei is
also more common. Most of the
localities in which T. chinensis is
the predominant species are in
Chongqing and adjacent W Hubei. In Viet Nam it has been found in Ha Giang province near the
Chinese border and further S in Hoa Binh and Son La provinces, where it occurs on karst lime-
stone ridges and summits. In most of China Taxus chinensis occurs in evergreen and deciduous
broad-leaved forests, frequently along streams. Its altitudinal range in China is from 1100 m to
2700 m a.s.l.; in Viet Nam between 900 m and 1500 m. Taxus chinensis is listed as Endangered (EN)
on the IUCN Red List due to continuous decline from deforestation and exploitation.

Map MAJ-124. Taxus mairei [n=82,

m=80, h=12, 1908–2011]

Taxus mairei has a similar distribu-

tion as T. chinensis (map MAJ-123)
but it is more common in the E
and extends to Taiwan. In the S it
does not reach Viet Nam and is rare
in Guangxi. In Sichuan it is wide-
spread and more common than T.
chinensis. As is also demonstrated
by the herbarium collection data
mapped in Ying et al. (2003: 192–
193) the province of Hunan appears to be nearly without Taxus and only in the far S of the prov-
ince in the Nan Li Shan near the border with Guangdong and in the far NE of the province a few
collections of T. mairei have been made. There are mountainous areas in this province in all but
the NE so it is not easily explained why Taxus should be absent there. Taxus mairei is common
as a shrub in more or less open vegetation, especially on limestone; in forests it is usually follow-
ing streams. Its altitude range is substantial, from 100 m a.s.l. in some localities in the E to >3000
m in the far W of its range as well as in Taiwan. This species is listed as Vulnerable (VU) on the
IUCN Red List.
252 mainland asia and japan

Taiwan

Taiwan is a mountainous island separated by the Taiwan Strait at a distance of ca. 180 km from main-
land China. It has a total land area of 35,883 km² and is 394 km long and 144 km wide. More than 30%
of the island rises above 1000 m and more than 200 peaks are over 3000 m high, with Yü Shan (Jade
Mountain) at 3952 m the highest mountain. There are coastal plains and hills in the W and S but on
the E side the mountains drop steeply to the Pacific Ocean in most places, indicating the geological
origin of the island as a tilted fault bloc. The climate is subtropical but temperate to cool temperate
in the central mountains; there is no permanent snow. The natural flora of the lowlands has strong
similarities with the nearby provinces of mainland China, but the montane forest flora has many
affinities with distant Yunnan and Sichuan, or even with the E Himalaya at alpine elevations. There
are 26 species of conifer in Taiwan in five families and 16 genera; eight species are endemic. Given its
limited area, slightly larger than the land area of the Netherlands, this is a remarkably high diversity.
There are taxonomic links with W-central China and the E Himalaya, but also with N Viet Nam and
adjacent parts of Lao PDR. Taiwania cryptomerioides (map MAJ-89 on p. 230) and Cunninghamia
konishii (map MAJ-88 on p. 230) are the best examples of this type of distribution. There are also links
with Japan, e.g. Chamaecyparis obtusa var. obtusa (map MAJ-183 on p. 288) but here separation has
led to two varieties, one of which is endemic to Taiwan and the other to Japan.

Map MAJ-125. The distribution of conifer species in Taiwan.

Conifers are with very few exceptions distributed in the moun-

tainous parts of Taiwan and are virtually absent in the plains
on the E side of the island and in the plain making up most of
Ilan County in the NE. However, they are also absent or rare
on the lower E flank of the Central Mountain Range and in the
Taitung Coastal Mountain Range. This side of the island receives
very high rainfall and the forests are dominated by angiosperms.
Most of the lowlands and foothills are deforested and in cultiva-
tion, which undoubtedly has an impact on the natural distribu-
tion of those conifer species that could occur there. But the great
majority of species are montane to high montane/subalpine and
so the distribution as shown is mostly natural. Note the occurrence of conifers (Podocarpus cos-
talis) on Orchid Island (Lan Yü) E of the southern tip of Taiwan. Below we present the eight
endemic species; all the other species are treated in other sections of the chapter Mainland Asia
and Japan and, in the case of P. costalis, in the chapter Malesia (map MA-14 on p. 302).

Pinaceae

Map MAJ-126. Pinus morrisonicola [n=38, m=22, h=7, 1898–2001]

Pinus morrisonicola is a montane pine with a scattered distribution

but with a main occurrence in two areas in the N half of the island.
A few localities are near the E coast; it appears to be absent in the S
part of Taiwan. Its altitude range is considerable, between 300 m and
2300 m, so its distribution is not so much limited by climate zones
but rather by site availability. Pinus morrisonicola occurs mostly on
steep rock outcrops and ridges, or occasionally it can invade land
slide sites where the forest has been removed. Such occurrences can
be ephemeral if forest succession leads to a return of more shade toler-
ant angiosperms.
 mainland asia and japan 253

Map MAJ-127. Pinus taiwanensis var. taiwanensis (black) [n=36, m=31, h=7,

1906–2005]; P. taiwanensis var. fragilissima (red triangle) [n=1, m=1, h=3,
1991–1991]

Pinus taiwanensis is the more common of the two endemic species

of Pinus in Taiwan and is distributed in the mountains in the central
part of the island, with only a few outliers near the E coast in the
Taitung Valley and in the far S of the island. In one of the southern
localities, in the Kuan Shan near the village of Wulu, P. taiwanensis
var. fragilissima occurs in the same area as var. taiwanensis. This vari-
ety (originally described as a new species) is still only known from its
type locality. In the mountains of the interior of Taiwan P. taiwan-
ensis occurs from 800 m to 3000 m a.s.l., with exceptions to 3400 m
where it is severely stunted. In the Taitung Valley it comes down to
600 m. It is often associated with various species of oak (Castanopsis,
Quercus); at higher altitudes also with Pinus morrisonicola and P. armandii var. mastersiana and
with Pseudotsuga sinensis.

Map MAJ-128. Picea morrisonicola [n=16, m=14, h=8, 1906–2002]

Picea morrisonicola is distributed in the Central Mountain Range

(Chungyang Shanmo) along the middle part of its length. This spe-
cies occurs S to 23° 13’ N, i.e. just S of the Tropic of Cancer, which
makes it the southernmost species of spruce in the world (P. farreri
in Myanmar [Burma] reaches 26° N, see map MAJ-138 on p. 259). This
spruce is a high montane species, occurring at elevations between
2300 m and 3000 m on N-facing mountain slopes and in ravines. The
soils are acid and podzolized. The climate is cool temperate, with
a monsoon character and very wet: annual precipitation exceeds
4000 mm. In some areas it forms almost pure stands, elsewhere it is
associated with other conifers and on lower slopes with angiosperms
such as Quercus (Cyclobalanopsis), Carpinus, Pittosporum and Ulmus.
At higher altitudes it can grow together with Abies kawakamii and
Juniperus squamata, but the subalpine conifer forests are Abies and Juniperus only and Picea stays
behind; the tree line, unlike in many other mountains, has no spruces. Logging has reduced the
area of occupancy (AOO) of this species which is listed as Vulnerable (VU) on the IUCN Red List.

Map MAJ-129. Abies kawakamii [n=30, m=25, h=9, 1906–2006]

Abies kawakamii is a high montane/subalpine species occurring

in the highest parts of the Central Mountain Range, at altitudes
between (2300–)2400–3800(–4000) m, i.e. up to the tree line. At
the lowest altitude it is a component of mixed forest with Quer-
cus spp., higher up soon joined by Tsuga chinensis, Pinus armandii,
Picea morrisonicola and other conifers. At ca. 3000 m it becomes the
dominant tree, associated with Juniperus squamata var. morrisoni-
cola or in pure stands and eventually at extreme altitude as dwarfed
krummholz. These stands of Abies are often surrounded by subal-
pine bamboo grassland. This bamboo (Yushania niitakayamensis)
also occurs as ground cover under the trees (see photo).
254 mainland asia and japan

Abies kawakamii in the Central Mountain Range, Taroko National Park, Taiwan. Photo credit Aljos Farjon.

Podocarpaceae

Map MAJ-130. Podocarpus nakaii [n=17, m=15, h=9, 1912–1995]

Podocarpus nakai has a scattered distribution in the N half of Tai-

wan and is perhaps most common around Sun-Moon Lake in Nan-
tou County, where most herbarium collections have been made.
An outlying population occurs in the far NE of the island in moun-
tains near Tawu. This population was still extant in 1976 when it
was sampled by C. N. Page for the Edinburgh herbarium. The most
western locality on the map is based on a collection made in 1912
(W. R. Price 1094 at K) “between Shan-hsi and Fan-po” in a forested
valley. It is likely that this population has disappeared in the cen-
tury since its discovery. The same may be true of the locality fur-
ther N on the coastal plain, the herbarium collection is from 1940.
Flora of Taiwan, ed. 2, 1: 565 (1994) only mentions Nantou County
for its distribution. Podocarpus nakai occurs as scattered individual trees in colline to lower mon-
tane forest dominated by Fagaceae, Lauraceae, Theaceae, Magnoliaceeae and other angiosperms.
Deforestation and general logging has greatly reduced the area of primary forest in which this
species grows; as a result it has declined rapidly in the past century and is now listed as Endan-
gered (EN) on the IUCN Red List.

Cupressaceae

Map MAJ-131. Calocedrus formosana [n=20, m=15, h=12, 1900–2002]

Calocedrus formosana occurs in the Central Mountain Range in the

N half of Taiwan. Its furthest S population is probably in the Sun-
Moon Lake area in central Nantou County. Early herbarium collec-
tions made in 1900 and 1918 in Pingtung County in the far S of the
islands came from planted trees and these have been excluded from
the map. The altitude range of this species is (300–)800–2000 m a.s.l.
Calocedrus formosana occurs in primary mixed forest where it can
be an emergent tree, but also in secondary forest and in more open
situations on rocky slopes and ridges. It has declined sharply in the
last century due to logging, so that these large emergent trees are now
rare and confined to national parks and other protected areas. Caloce-
drus formosana is listed as Endangered (EN) on the IUCN Red List.
 mainland asia and japan 255

Map MAJ-132. Chamaecyparis formosensis [n=37, m=34, h=13, 1896–2001]

Chamaecyparis formosensis occurs in the Central Mountains from S

Taipei County to N Pingtung County; a disjunct population is present
on Tai-he Shan in the far S of Taiwan. This species grows in the cool
temperate coniferous forest belt between 1700 m and 2900 m a.s.l.,
with Chamaecyparis obtusa var. formosana and C. formosensis as the
dominant species, associated with Calocedrus formosana, Cunningha-
mia konishii and Taiwania cryptomerioides, in the upper part of the
zone also Tsuga chinensis; in the under-storey occur Photinia davidi-
ana var. niitakayamensis and the bamboo Yushania niitakayamensis,
but often little else. Chamaecyparis formosensis is ­exceptionally long-
lived, estimated in excess of 3000 years, and can attain great sizes;
several of these giant old trees are now tourist attractions. This great
tree has been heavily exploited and few substantial stands remain, now mostly in national parks
and other reserves. The conversion of these forests into plantations with the Japanese conifer Cryp-
tomeria japonica has been the main threat and has not entirely ceased. Increased risk of forest fires
by opening up the forests with logging roads is another threat to this very slow growing conifer.
Chamaecyparis formosensis is listed as Endangered (EN) on the IUCN Red List.

Chamaecyparis formosensis, C. obtusa var. formosana and (uncommon) Taiwania cryptomerioides in pri-
mary forest on Chilan Shan, Ilan County, Taiwan. The undergrowth is made up of saplings of the two spe-
cies of Chamaecyparis, the bamboo Yushania niitakayamensis and a thick carpet of mosses. Photo credit
Aljos Farjon.

Taxaceae

Map MAJ-133. Amentotaxus formosana [n=14, m=10, h=7, 1925–1999]

Amentotaxus formosana is very rare and confined to the southern-

most part of Taiwan, occurring in the low mountains that form an
extension of the Central Mountain Range. Not many herbarium col-
lections have been made since it was first collected in 1925. This
tree is often difficult to find in dense evergreen subtropical forest,
occurring under canopy of angiosperm trees amid tree ferns, ferns,
shrubs and tall sedges. Its altitudinal range is between 500 m and
1300 m with most trees occurring above 900 m a.s.l. The natural for-
est in which this species occurs is under threat from conversion, at
the lowest altitude to palm plantations (Areca catechu) and higher
up to conifer plantations with Cryptomeria japonica. Amentotaxus
formosana is listed as Endangered (EN) on the IUCN Red List.
256 mainland asia and japan

Indochina

Geography

Indochina is defined in this chapter primarily as a biogeographical region (the Indochinese

Region) but with boundaries of nation states. It is the region between India, China and Malesia;
the latter includes Peninsular Malaysia and is the biogeographical region of a separate chapter
Malesia in the Atlas. The following countries are here included: Cambodia, Lao PDR, Myanmar
[Burma], Thailand and Viet Nam. The region lies almost entirely within the N tropical latitudes
with the exception of N Myanmar and northernmost Viet Nam. It is a large peninsula surrounded
by the Indian Ocean in the W and the South China Sea in the E, with to the S an extended
and much narrower peninsula which includes parts of Myanmar, Thailand and all of Peninsular
Malaysia. Indochina is mostly mountainous in the N half and on the E side along the border
between Lao PDR and Viet Nam (Annam Highlands), but with intervening lowlands. The S half is
predominantly lowland except in Viet Nam. Many large rivers flow generally from N to S through
it, some like the Irrawaddy in Myanmar and the Mekong in Viet Nam forming huge deltas extend-
ing into the ocean. In particular the lowlands and broad river valleys are densely populated and
have mostly been under agriculture for several thousand years. There are obvious biogeographical
connections (the Malesian Region has already been mentioned) with other areas, of which the
most important in terms of conifer distribution is Yunnan in S China. Yet the region is sufficiently
distinct to contain several endemic or near endemic species.

Pinaceae

Map MAJ-134. Pinus latteri [n=47,

m=43, h=12, 1909–2007]

Pinus latteri is closely related to

P. merkusii (map MA-74 on p. 338
and is perhaps only a subspecies,
yet its distribution is separated as
it occurs in mainland SE Asia only.
It is present in all five countries
and in China, from SE Myanmar
[Burma] to Guangdong (Leizhou
Peninsula) and Hainan Island in
S China and to S Viet Nam. The
presence in Hainan is perhaps
due to an early introduction
and its status there is in need of
verification; only two herbarium
collections were found and one
could not be located. Pinus latteri
occurs from near sea level to ca.
1200 m a.s.l., forming more or less open stands on old river terraces with sandy or gravelly soil or
in seasonally dry hills. It is a fire-adapted species capable of surviving grass fires through a ‘grass
stage’ as a seedling and can quickly invade open terrain. Its range is within the influence of SE
monsoons, with mean annual precipitation around 1500 mm in Myanmar and Thailand. It is also
a constituent tree in dipterocarp forests (e.g. Dipterocarpus tuberculatus), especially in more open
situations on drier sites of ridges in the hill country of Indochina.
 mainland asia and japan 257

Pinus latteri in Mae Sarin National Park, Thailand. Photo credit H. Hazebroek.

Map MAJ-135. Pinus kesiya

var. kesiya (black) [n=42,
m=39, h=8, 1850–2004];
P. kesiya var. langbianensis
(red) [n=51, m=49, h=17,
1841–2008]

Pinus kesiya is the most

widespread and com-
mon species of pine in
SE Asia. The two vari-
eties are not sympat-
ric even though their
extent of occurrence
(EOO) does overlap in
part. Pinus kesiya var. kesiya has a more W distribution and is restricted to mainland SE Asia,
where it is present in the Khasi Hills, the Naga Hills and Manipur in NE India, in Myanmar [Burma]
and Thailand, in the Tumpor Mountains of W Cambodia, in Lao PDR and in various localities in
Yunnan, China. Pinus kesiya var. langbianensis occurs in the most S part of Yunnan bordering on
Myanmar and Lao PDR as well as in Malipo Xian near the Viet Nam border and just S of it in Viet
Nam; also in Saravane Province in S Lao PDR and in S Viet Nam, where a major population is in
the Lang Bian Mountains in Lam Dong Province. Finally it is present in extensive stands in the
Cordillera Central and Zambales Mountains of N Luzon, Philippines. Pinus kesiya occurs in pine
savannas, pure stands with nearly closed canopy, and mixed pine-angiosperm forests in valleys
with e.g. Quercus serrata and Alnus nepalensis along streams. It occupies drier sites in NE India,
Myanmar and Thailand, at altitudes generally between 800 and 1500 m a.s.l., occasionally to 2000
m. Further E in Lao PDR, Viet Nam and on the island of Luzon in the Philippines, its altitudinal
range is greater and it is found up to 2700–3000 m a.s.l. in a much wetter climate. Pinus kesiya is a
tropical pine which does not survive frost; in China it is replaced by P. yunnanensis (map MAJ-43
on p. 205) further N in Yunnan and Sichuan.
258 mainland asia and japan

Map MAJ-136. Pinus krempfii [n=12, m=12, h=8,

1900–2006]

Pinus krempfii is endemic to Viet Nam and

has a limited distribution in the mountains
N and NE of the city of Da Lat. The most
extensive populations are in the Bi Doup
Massif and the Hon Vong Phu Massif;
Chu Bao Mountain, now a forest reserve
where the type collection was made by
M. Krempf, is the farthest NE locality. This
species, although now classified within
subgenus Strobus, is morphologically dis-
tinct from all other pines, most notably in
its broad, flat leaves. Pinus krempfii grows
in primary closed evergreen tropical mon-
tane rainforest, where it is an emergent
above the general canopy dominated by
angiosperm trees. This habitat and place
in the forest succession is unusual for a
pine and its broad leaves may be an adap-
tation to compete for light with angiosperms. Too little is known about the ecology of this species
to pronounce this to be the case, but comparison with other emergent conifers in tropical forests
(e.g. Dacrydium xanthandrum, map MA-32 on p. 310) and a study of their role in the forest, as
was done for some New Zealand species (Enright & Hill, eds. 1995) is to be recommended. Its
altitudinal range is between 1200 m and 2000 m a.s.l. and it is often found on steep slopes and on
mountain ridges. Although several populations are now in protected areas, logging has caused its
disappearance from other localities and P. krempfii is therefore listed as Vulnerable (VU) on the
IUCN Red List.

Pinus krempfii emergent trees in tropical montane evergreen angiosperm rainforest at Doc Cong Troi, Lam
Dong Province, Viet Nam. Photo credit Philip Thomas.
 mainland asia and japan 259

Map MAJ-137. Pinus dalatensis [n=30, m=30, h=8, 1967–2011]

Pinus dalatensis is distributed in the Annam Highlands,

mostly in Viet Nam but known from at least one locality
in Lao PDR (Nakai Nam Theun NBCA, Phou Ac escarp-
ment in Khammouan) with a population on both sides
of the border. This is the northernmost location of this
species; the southernmost localities are in the Bi Doup
Massif near the city of Da Lat. Pinus dalatensis is a rare
species occurring in mostly small and isolated popula-
tions, some of which show small morphological differ-
ences from other populations and have been described
as distinct taxa. It is a montane pine occurring at alti-
tudes between 1400 m and 2300 m in tropical evergreen
forest. This species is probably dependent on occasional
fires for regeneration (removal of competing vegetation),
but increased fire frequency caused by people benefits
P. kesiya rather than this more slow growing species. Protec-
tion in several national parks and other reserves has gone
some way to prevent decline, but this decline continues in
many non-protected areas of forest. Increased activity in
forest surveys has discovered many small populations
in recent years, but this work has only just begun to reveal
the occurrence of this species in the Lao PDR and more
localities can be expected to be found there. The IUCN Red
List gives this species the status of Vulnerable (VU).

Map MAJ-138. Picea farreri [n=6, m=6, h=2, 1924–2002]

Picea farreri is a very rare and localized species

which occurs in mountains between the Nu Jiang
(Salween River) and N’maikha River along the bor-
der between China and Myanmar [Burma]. George
Forrest was the first ‘plant hunter’ to collect it in
Myanmar probably on Mt. Chimili in 1924 (G. Forrest
24828 at E) but the species was recognized as dis-
tinct from specimens collected by Reginald Farrer in
1932 in the upper Feng Shui Ling (valley) leading to
the pass into China. It also occurs on the pass (2800
m) and down the Chinese side. Since the 1930s no
further collections were made, in part due to politi-
cal circumstances, and it was thought that Picea far-
reri was limited to a very small area. New botanical
surveys by teams of botanists from China and the
West collected specimens in 1996 and 2002 in two locations further N in mountains between the
Nu Jiang and the Myanmar border, in Nujiang Lisu Prefercture and near Gongshan (Gaoligong
Shan), the farthest N so far. An absence of seed cones with these later specimens makes identi-
fication less than 100% certain, but this species is inferred on biogeographical grounds. It would
appear then that this rare spruce occurs in scattered localities along ca. 250 km of the mountain
chain that forms the border between the two countries. Picea farreri is listed as Vulnerable (VU)
on the IUCN Red List, downlisted from Endangered (EN) due to new discoveries in China.
260 mainland asia and japan

Podocarpaceae

Map MAJ-139. Nageia fleuryi [n=31,

m=31, h=8, 1918–2006]

Nageia fleuryi is the only species

in the family Podocarpaceae with
a distribution that is mostly in
Indochina. Other species, some
common and widespread in the
region, have their main distribu-
tions in Malesia and are treated in
that chapter. Nageia fleuryi occurs
in Viet Nam from the Chinese bor-
der S to Mt. Ba Na in Da Nang and
in S China in Guangdong, Guangxi
and Yunnan, mostly in the S parts
of these provinces and very scat-
tered. Only one recent (2006) her-
barium record has been obtained from Lao PDR, collected at the Tad Leuk waterfall in Khammouan
(M. F. Newman et al. LAO 1534 at E). With botanical surveys increasing in that country it is likely
to turn up elsewhere, but it seems a rare species and is replaced in the W and S by N. wallichiana
(map MA-55 on p. 323) with which it is sometimes confused. Nageia fleuryi is most often found
on karst limestone formations in mixed angiosperm/conifer forest, which explains its predomi-
nance in N Viet Nam and S China. It is less common in tropical rainforest dominated by evergreen
broad-leaved angiosperm trees, which tend to develop on richer substrates. It is a (sub)montane
species, but its altitude range (in Viet Nam) is somewhat broadly defined as between 500 m and
1200 m.

Cupressaceae

Map MAJ-140. Xanthocyparis vietnamensis [n=9,

m=9, h=6, 1999–2012]

Xanthocyparis vietnamensis was discovered in

the late 1990s near Bat Dai Son, in the karst lime-
stone formations along the Chinese border in Ha
Giang Province, Viet Nam. Subsequent surveys
have found it in nearby localities, e.g. Sing Xuoi
Ho and it is now known in this province from
several localities, all in Quan Ba District. In 2012
a tree was discovered in Mulun Nature Reserve,
Huanjiang Xian, Guangxi Province, China and
herbarium specimens collected from it are in
IBK (Y. S. Huang et al. 1432–1433, coll. 27–4–2012).
Xanthocyparis vietnamensis occurs, with other
conifers and small-leaved angiosperm trees and shrubs, on karst limestone mountains which cover
vast areas in this region. The locality in China, ca. 450 km NE of the Vietnamese population, seems
oddly distant given that such karst mountains occur in much of the intervening country and indeed
are continuous across the international border, from which Bat Dai Son is only a few km away.
 mainland asia and japan 261

This extremely rugged terrain is difficult to explore; the karst mountains can have perpendicular
flanks and knife-edge ridges covered in dense vegetation. The conifers are on the ridges and summit
crests avoiding competition from large leaved evergreen angiosperms which occupy the talus slopes
below. Exploring these numerous tower mountains extensively will be a major undertaking involv-
ing many more trained botanists than are currently available. As our knowledge stands at present,
evidence of exploitation and low numbers of mature trees in all known localities (only one tree
found in China so far!) have put this species as Endangered (EN) on the IUCN Red List.

Habitat of Xanthocyparis vietnamensis near Bat Dai Son, Ha Giang Xanthocyparis vietnamensis in
Province, Viet Nam. Photo credit Aljos Farjon. Mulun N.R., Guangxi, China.
Photo credit Y. S. Huang.

Map MAJ-141. Calocedrus rupestris

[n=16, m=16, h=7, 1999–2009]

Calocedrus rupestris was discov-

ered in 1999 in Cao Bang Prov-
ince in N Viet Nam, close to the
border with Guangxi Province,
China. Subsequent collections
made in botanical surveys estab-
lished its distinction as a species
in other locations in N Viet Nam
and in 2008–2009 it was found in
three localities in Guangxi. It will
undoubtedly be found in other
localities in this region, possibly in
SE Yunnan along the border with
Viet Nam where extensive areas
have similar habitat. In Viet Nam
it has so far been found in seven
provinces N of the 17th parallel.
Calocedrus rupestris is another
conifer species occurring on the
karst limestone mountains that
are a dominant geological feature
in this region. The result of deep
262 mainland asia and japan

erosion of a vast plateau of Palaeozoic limestone, these very steep mountains and towers have a
distinctive flora characterized by more than a dozen species of conifer (not all endemic to these
formations), small leaved angiosperm trees and shrubs, epilithic orchids, ferns and mosses. Soil is
almost absent and although rainfall is high, much of it runs off immediately. Most of the forests in
the intervening valleys have been cut and replaced by various forms of agriculture; where pockets
remain on talus slopes or in deep recesses it is large leaved evergreen tropical rainforest in which
none of the conifers present on the mountains could compete and survive. Inaccessibility is now
also their best survival strategy against human exploitation, albeit not full proof. Although only
known by botanists for a short time, evidence of exploitation suggests a continuing decline and
C. rupestris is listed as Endangered (EN) on the IUCN Red List.

Cephalotaxaceae

Map MAJ-142. Cephalot-
axus mannii [n=56, m=54,
h=10, 1836–2006]

Cephalotaxus mannii is
widely distributed in
Indochina and occurs
from Megalaya, Nagaland
and Arunachal Pradesh
in NE India and the Yar-
lung Zangbo-Brahma-
putra Bend in Xizang
[Tibet] to Hainan Island,
Guangxi and W Guang-
dong in S China and the
mountains N of Dalat
in Lam Dong Province,
Viet Nam. This species
is probably widespread
in N Myanmar [Burma]
but recent collections
(post 1962) are not available; it is also fairly common in NW Thailand and in N Viet Nam and
occurs across the Vietnamese border in SE Yunnan, China. It is reported from Lao PDR but we
have no herbarium records to show its localities in that country. This species is present in small
numbers in mixed evergreen or (semi-)deciduous forests and can become a tall tree competing
with angiosperms in the canopy. Its altitude range is from 500 m to 2000 m a.s.l. Deforestation in
Indochina and China and logging of the larger trees are causing continuous decline and despite
its wide distribution this species is listed as Vulnerable (VU) on the IUCN Red List.
 mainland asia and japan 263

Taxaceae

Map MAJ-143. Amentotaxus hatuyenensis

(black) [n=1, m=1, h=1, 1979]; A. poilanei (red)
[n=2, m=2, h=6, 1946–1995]

These two species remain poorly known

to science. Amentotaxus hatuyenensis
is known from the type collection (Vu
Xuan Phuong 329 at P) made in August
1979 near Lung Cun on the border with
China, Ha Tuyên Province, and reported
from Ha Giang Province, both in N Viet
Nam. Of A. poilanei two herbarium col-
lections are known from one locality,
first the type E. Poilane 32686 (A, BM,
P, US) in December 1946 and subse-
quently L.V. Averyanov et al. VH-774
in March 1995 (LE, MO). The local-
ity of this species is in the Ngoc Linh
Mountains in Gia Lai Province, central
Viet Nam. Amentotaxus hatuyenensis is
another rare species of conifer occur-
ring on the karst limestone mountains
of N Viet Nam and possibly across the
border in China. Since there are two
other species native to this region, A. argotaenia (map MAJ-116 on p. 246) and A. yunnanensis
(map MAJ-117 on p. 247) and especially juvenile plants are quite similar, only a critical revision
of available and new material, which would have to include DNA comparison, can give more
insight in the taxonomic status and distribution of A. hatuyenensis. Amentotaxus poilanei is appar-
ently a large tree occurring in high montane closed evergreen rainforest, at an altitude around
2300 m a.s.l. It is locally common but scattered, mixed with angiosperm trees and perhaps Nageia
­wallichiana as the only other conifer present. Whether it will be found in other localities remains
to be seen. The fact that no second collection was made in 50 years and then only from the same
locality indicates that it may be an extremely local endemic. Both species are in a category of
threat on the IUCN Red List (EN and VU respectively) mainly due to perceived rarity and decline
of potential habitat.
264 mainland asia and japan

Mainland Asia Far East

Geography

In this Atlas the Far East of mainland Asia includes the Russian Far East, i.e. Primoriye, Sakhalin
Island, the Kuril Islands, mainland regions bordering the Sea of Okhotsk, Kamchatka and the
farthest E of Siberia up to the Bering Strait, and Korea (North Korea and South Korea). With the
exception of Korea, this region is very thinly populated resulting in mostly undisturbed forests
and landscapes. On the mainland, these form a continuation of E Siberia with the exception of
Primoriye bordering on the warmer Sea of Japan. Its Sikhote Alin Range has a moderate elevation
to ca. 2000 m and a more diverse forest cover than the Siberian taiga and Kamchatka. Korea has
a varied topography and with Cheju-do (Quelpart Island) extends S to ca. 33° 24’ N and into a
mild maritime climate. Sakhalin is a 948 km long island N of Hokkaido in the Sea of Okhotsk near
the mainland coast, with low mountains to ca. 1600 m and a cold maritime climate; it is mostly
covered with coniferous forests. Kamchatka is a ca. 1400 km long peninsula connecting with NE
Siberia and bordered in the W by the Sea of Okhotsk and in the E by the Pacific Ocean and the
Bering Sea. It has several mountain ranges and numerous volcanoes, the tallest of which is Sopka
Klyuchevskaya at 4750 m. Its only conifer forest is a taiga remnant (from climate oscillations,
not human interference) in the central Kamchatka Valley. The Kuril Islands form a connecting
string of islands between Kamchatka and Hokkaido (Japan); only the southernmost islands are
partly forested. There is considerable overlap with other areas in Asia in the ranges of most of the
conifer species that occur here, yet the region has a few endemic taxa, including a monospecific
genus. Only endemic species and those with their major distribution in Mainland Asia Far East
are mapped and discussed here.

Pinaceae

Map MAJ-144. Pinus pumila

[n=67, m=64, h=16, 1784–2007]

Pinus pumila is widespread

in E Siberia, Kamchatka
including Bering Island (the
westernmost island of the
Aleutian chain), the N Kuril
Islands, Sakhalin Island,
Primoriye, Japan (Hok-
kaido and Honshu), Korea,
NE China and NE Mongo-
lia. This dwarf pine forms
extensive low thickets, in
the NE of its range in lowlands, but elsewhere mostly in mountains and in Korea and Honshu on
or near mountain summits. In Japan the Dwarf Siberian pine occurs from 1400 m to 3200 m a.s.l.,
but on the Kamchatka Peninsula it is found from sea level up to 1200 m in favorable localities.
It can also occur in the under-storey of taller conifer forests, mostly on sites with shallow, stony
soil or peat.
 mainland asia and japan 265

Pinus pumila (large dark green

patches) in central Hokkaido,
Japan at ca. 1500 m. Photo
credit and © Dendrological
Atlas Team (Zsolt Debreczy
& István Rácz).

Map MAJ-145. Pinus koraiensis

[n=47, m=42, h=16, 1842–2004]

Pinus koraiensis occurs in

regions surrounding the
Sea of Japan. In China it is
found in Heilongjiang, Jilin
(mainly in the Shangbai
Shan in the SE) and E Lia-
oning. In Russia it occurs
in Primoriye. Through the
Korean Peninsula this spe-
cies is common in North
Korea (but we could not
access enough herbarium
locality data to show this)
and along the central
mountains of South Korea.
In Japan it is common in
the central mountains (Jap-
anese Alps) of Honshu but uncommon further N, extending as far as Hokkaido. The farthest from
the sea are some populations in Heilongjiang. In Primoriye this pine grows from 200 m to 600
m a.s.l., from 500 m to 1300 m in China, and in Japan it occurs to an altitude of 2500 m. Pinus
koraiensis is present in deciduous angiosperm forest as well as in mixed coniferous forest, with a
shift from the former to the latter with increasing altitude and latitude. Exploitation in Korea has
led to decline, but in many parts of its range it is still common.
266 mainland asia and japan

Map MAJ-146. Larix gmelinii var. gmelinii (black) [n=40, m=39, h=7, 1844–2003]; L. gmelinii var. japonica
(red) [n=17, m=15, h=9, 1861–2003]; L. gmelinii var. olgensis (yellow) [n=30, m=29, h=8, 1860–1986]; L. gme-
linii var. principis-rupprechtii (blue) [n=25, m=25, h=5, 1909–1996]

Larix gmelinii is a highly variable species with a wide distribution in the NE parts of Asia. Four
distinct varieties are recognized and these occupy different ranges. The most widespread is
L. gemilinii var gmelinii, a Siberian species occurring from Lake Baikal and the Jenissei River in
Siberia eastwards to the Sea of Okhotsk and the Bering Sea. The map for this taxon is incomplete
due to a lack of sufficient herbarium specimens in the Conifer Database, but it includes the dis-
junct population in Kamchatka (which Russian botanists often consider to be a separate species).
Larix gmelinii var. japonica occurs in Hokkaido (Japan), Sakhalin and the southernmost Kuril
Islands (Russia). In Primoriye (Russia), Heilongjiang and Jilin (China) and North Korea occurs
L. gmelinii var. olgensis, while L. gmelinii var. principis-rupprechtii is endemic to China with a
distribution in the mountains that form the dividing highlands between the Northern Plain and
the steppes and deserts of Nei Monggol (Inner Mongolia). The Siberian variety is a constituent
of ‘light taiga’ where fires are most frequent; this variety is also extremely hardy and forms the
tree line towards the Arctic Ocean, where dwarfed trees that escaped fires can be hundreds of
years old. The other varieties, occurring further S and under the maritime influence of the North
Pacific, are part of the montane mixed conifer forests of the Russian Far East, former Manchuria
and Hokkaido. Larix gmelinii var. principis-rupprechtii attains the highest altitude and has been
found at 3000 m a.s.l. but the maximum altitude for most varieties is around 2000 m.

Map MAJ-147. Abies koreana [n=16, m=16, h=5, 1907–1996]

Abies koreana is a rare species with a scattered dis-

tribution in North Korea and in South Korea. Only
one herbarium collection in the Conifer Database is
from North Korea; it was collected on Mt. Daeseong
in 1996. Most collections are from Halla-san on Cheju
(Quelpart) Island; the remaining records are from
Chiri-san, Ungseokbong Peak and Tokyu-san, all in
mainland South Korea. Other known localities there
are Kaya-san, Kongo-san and Mudung-san; for lack of
available collections these are not shown on the map.
This species forms small and isolated populations on
or near the summits of mountains between 1000 m
and 1900 m altitude. Abies koreana grows in pure
stands or mixed with Betula ermanii on Cheju Island;
 mainland asia and japan 267

on the mainland it is also mixed with Picea jezoensis, Pinus koraiensis, Taxus cuspidata, Quercus
mongolica var. mandshurica, Cornus controversa, Acer spp., and several genera of low shrubs, e.g.
Juniperus, Deutzia, Ribes and Rhododendron. Abies koreana is listed as Endangered (EN) on the
IUCN Red List, while it is common and widespread in horticulture.

Cupressaceae

Map MAJ-148. Microbiota decussata [n=14, m=13, h=11,

1921–1994]

Microbiota decussata is the only instance of a mono-

specific conifer genus in Asia outside China and Japan.
The distribution of this decumbent shrub is limited to
the Russian Far East and concentrated in the S part of
Primoriye, E of Vladivostok at the end of the Sikhote
Alin Range. Outside this area it has been collected in
a few isolated localities, the northernmost of these
is on the Anyuy River near the confluence with the
Amur River. Russian Floras indicate other locations
in the Sikhote Alin Range but we had no herbarium
specimens available to map these. Microbiota decus-
sata is a decumbent shrub in montane habitats both in and outside forests, at altitudes between
800 m and 1200 m a.s.l. It occurs in mixed conifer or angiosperm-conifer forest with e.g. Abies
nephrolepis, Picea jezoensis, Pinus koraiensis, Acer ukurunduense, Alnus maximowiczii, Betula
ermanii, and Sorbus amurensis. In subalpine non-forest vegetation it can be assosiated with Pinus
pumila, Juniperus sabina var. davurica, or Rhododendron mucronulatum, often growing on talus
slopes or mossy granitic boulders with small accumulations of soil. It is listed as rare in the Red
Book of Russia and it is included in Nedoluzhko’s (1999) Endangered woody plants of the Rus-
sian Far East. The Conifer Specialist Group of IUCN was not able to place it in any category
of threat.

Map MAJ-149. Thuja koraiensis [n=23, m=22, h=9,

1897–2010]

Thuja koraiensis is a near endemic of the Korean Pen-

insula but it also occurs in the Changbai Shan in Jilin
Province, China. On the peninsula it is found in North
Korea (Pyongannam-do, Pyonganpuk-do, Hamkyong-
nam-do, Kangwon-do) and in South Korea (Kyongsan-
puk-do, Kangwon-do, Cholla-namdo). Thuja koraiensis
occurs on middle and upper slopes of mountains, at
altitudes between 750 m and 1950 m a.s.l. This species
is rare and scattered and in Korea the populations are
often very small and in many localities consist of low
shrubs or at most small trees. Yet in cultivation T. koraiensis appears to grow to a sizable tree; it
appears therefore that such trees have disappeared from the natural habitat. When E. H. Wilson
visited Korea in 1917–18 he saw no large trees, so it seems that these had mostly gone more than
a century ago. This seems to be the fate of many cupressaceous trees in East Asia, where their
wood has been valued above any other from conifers. This species is listed as Vulnerable (VU)
on the IUCN Red List.
268 mainland asia and japan

Japan

Geography

Japan is an archipelago of four large and many small islands situated on the easternmost and
westernmost edges respectively of the Eurasian and North American tectonic plates, separated
from mainland Asia by the Sea of Japan and the East China Sea. To the E is the Pacific Ocean,
where the Pacific and Philippine oceanic tectonic plates are subducted under the Eurasian and
North American plates in two of the longest and deepest ocean trenches in the world. This activ-
ity causes volcanism and uplift across Japan, making it one of the most active geological zones
around the Pacific. The main archipelago of Japan extends from Cape Soya-misaki in Hokkaido
at 45° 31’ N to the island of Yakushima at 30° 20’ N; to the SW the string of islands known as the
Ryukyu Islands or Nansei-shoto in Japanese reaches almost to Taiwan with Yonaguni Island at
24° 27’ S. Yet another string of islands extends far into the Pacific to the S of Honshu, of which
the Bonin Islands (Ogasawara-shoto) at ca. 27° N are relevant to conifer distribution. [Sakhalin
and the Kuril Islands, formerly Japanese, now belong to Russia.] Japan therefore reaches from the
cold temperate to subtropical climate zones. The largest islands of Japan, with Honshu (1300 km
long, 227,960 km²), Hokkaido to its N (77,980 km²), Kyushu to its SW (35,640 km²) and Shikoku
to its S (18,800 km²) are all mountainous, culminating in the famous volcano Fuji-san in Honshu
at 3776 m. Lowlands on the islands are mostly urbanized and the locations of the largest cities
as well as intensive agriculture. Some smaller satellite islands are also mountainous; of particular
significance for conifers is Yakushima off the southern tip of Kyushu.

Conifers of Japan

Families Genera Species Species + infra Countries with Area of occupancy Endemic taxa
conifers in km²
6/8 18/70 40/615 53/794 1/1 13,675/374,744 30
Numbers after ‘/’ are global or country totals.

Map MAJ-150. The distribution of coni-

fer species in Japan.

Conifers are present on nearly all

the islands of Japan, including many
of the remote oceanic islands in
the Ryukyu and Bonin archipela-
gos. The highest concentration of
conifer species is in the mountain-
ous central part of Honshu centring
around 36° N and 138° E and gradu-
ally diminishes towards both ends of
the island, becoming least in Hok-
kaido to the N and in Kyushu to the
S of Honshu. In Hokkaido exten-
sive coniferous forests occur but
with few species and here the map
shows a negative collection bias. In
Kyushu the warm temperate climate
 mainland asia and japan 269

tends to favour angiosperm dominated forests. More interestingly, there are some mountain-
ous areas in central Honshu with few or no conifer species, e.g. Fukui and Gifu Prefectures and
further W Hiroshima Prefecture, but on the whole the empty areas in Honshu are lowlands.
The outermost islands have only one species of Pinus, one species of Nageia, one of Podocar-
pus and two species of Juniperus. The diversity of conifer taxa in Japan is relatively high with
40 species, but biased towards taxa at higher ranks, with 18 genera and six families. One fam-
ily, Sciadopityaceae, is monospecific with Sciadopitys verticillata endemic to Japan. Cryptomeria
japonica (Cupressaceae) is also endemic, but it was introduced in China centuries ago and has
naturalized in some areas there. Thujopsis dolabrata (Cupressaceae) is endemic and restricted to
N Japan. Pinaceae is the most diverse family with 24 species (8 in Pinus, 6 in Picea, 5 in Abies).
Including subspecies and varieties, the total of taxa in Japan is 53 and 30 of these are endemic,
i.e. 56.6% is limited to these islands, many with restricted distributions but some occurring more
widely in Honshu or beyond.
The diversity maps MAJ-151–152 show that generic diversity is more widespread than specific
diversity. Apart from Honshu, Shikoku and Kyushu also have a few cells with more than five
conifer genera, but these islands do not reach the numbers of species attained in the (central)
mountains of Honshu. The distribution of narrow endemics on map MAJ-153 reflects this pattern
of diversity, but in addition the islands of Yakushima and Tanegashima have an endemic species,
Pinus amamiana. Highest values of local endemism are concentrated in two 0.5 dgree cells in
the S part of Nagano Prefecture, where Picea koyamae, Picea maximowiczii (2 varieties) and Picea
alcoquiana var. acicularis occur in an area with the highest conifer diversity of Japan, the Akaishi
and Yatsugadake Mountains.

Map MAJ-151. Map of Japan show-

ing numbers of conifer genera per 0.5
degree cell.

�‒�
�‒�
�‒�
�‒��
��‒��
270 mainland asia and japan

Map MAJ-152. Map of Japan showing

numbers of conifer species + infra-
specific taxa per 0.5 degree cell.

�‒�
�‒�
��‒��
��‒��
��‒��

Map-MAJ-153. Rarity values per 0.5

degree cell. Cell score is the total of
the rarity calculations for each spe-
cies in the cell. The rarity calculation
is1 divided by the total of cells occu-
pied by a species. Thus, a species
that occurs in one cell scores 1 while
a species that occurs in 10 cells scores
only 0.1.

Pinaceae

Of 11 genera in this family, six are present in Japan: Abies, Larix, Picea, Pinus, Pseudotsuga and
Tsuga. All are widespread elsewhere in the Northern Hemisphere, but of 24 species 14 are endemic
to Japan. Some others only extend to nearby Korea, a peninsula with a flora with many similari-
ties to Japan. These facts indicate that the isolation of Japan in biogeographical terms is relatively
recent for this family as it still shares 10 species with mainland Asia (compare New Caledonia and
New Zealand with similar numbers of conifer species but no sharing of species with anywhere
else, or Tasmania, close to Australia but with only two species of conifer in common).
 mainland asia and japan 271

Pinus

There occur seven species of Pinus in Japan, only two of these are endemic and two other species
extend only to South Korea. The other species have wider distributions. Of these, Pinus koraiensis
and P. pumila are excluded from this part of the Atlas on Japanese conifers because Japan con-
stitutes only a small section of their total ranges. Their maps are MAJ-145 on p. 265 and MAJ-144
on p. 264.

Map MAJ-154. Pinus thunbergii

[n=56, m=51, h=13, 1859–2003]

Pinus thunbergii has a wide but

scattered distribution in Japan
and Korea, where it mostly occurs
in lowland coastal localities and
on small islands. In Korea it is
only known to occur naturally on
islands, some well off shore. In
some localities, e.g. near Hitoana
in Shizuoka Prefecture, Honshu,
it was found high up at 900 m,
while on Tanegashima Island SE
of Kyushu it has been reported
from near sea level to 600 m alti-
tude. More commonly P. thunber-
gii is a pine that forms open forest or isolated stands on old beaches and stabilized dunes; in
some lowland areas e.g. near Tokyo such stands, if natural and not from plantations, are remnants
of perhaps more extensive pine forests. As it is a pioneer species, spread from plantations does
occur, and these may be in areas such as greater Tokyo where the species is also native, making
a detailed mapping of its natural distribution now almost impossible.

Map MAJ-155. Pinus den-

siflora [n=116, m=110, h=19,
1861–2001]

Pinus densiflora is distrib-

uted in lands around the
Sea of Japan and the Yel-
low Sea (Huang Hai) and
occurs in Japan (Honshu,
Kyushu, Shikoku), Korea,
China (Heilongjiang, Jil-
ing, Liaoning, Shandong,
Jiangsu, Anhui, Henan)
and Russia (Primoriye).
Pinus densiflora occurs
in extensive pure stands in many parts of its range and is one of the most dominant conifers in
Japan and Korea. It grows in a variety of acidic soils, from dry sandy or rocky sites to peaty soils. In
Japan it reaches from near sea level (and close to the shore) up to 2300 m in the mountains, but
on mainland Asia its altitudinal range is more restricted and extends upward to only 900 m in NE
China and 1300 m in Korea. In areas where broad-leaved forest dominates, P. densiflora is restricted
272 mainland asia and japan

to poorer sites such as rock outcrops on S-facing slopes and edges of moors or mountain lakes.
Pinus densiflora is closely related to P. sylvestris (Leslie et al., 2012; Parks et al., 2012) and may have
diverged as a result of climatic changes in the Pleistocene; their ranges do not overlap at present
(for P. sylvestris see map EM-5 on p. 143).

Pinus densiflora in S Kyushu,

Japan. Photo credit and ©
Dendrological Atlas Team
(Zsolt Debreczy & István
Rácz).

Map MAJ-156. Pinus luchuensis [n=24, m=24,

h=10, 1912–2005]

Pinus luchuensis is a ‘maritime’ pine

endemic to the Ryukyu Islands (Nansei-
shoto) where it occurs on many islands
(our data do not cover all with herbarium
specimens) in from N to S the Tokara
Group, the Amami Group, the Okinawa
Group and the Sakishima Group. These
islands form an arc around the southern
limits of the East China Sea between
Yakushima in the NE and Taiwan in the
SW and are of volcanic origin, arising
on the NW margin of a deep oceanic
trench. Hence this species of Pinus has
its origin in one or more dispersal events
from Japan, Taiwan or the Asian main-
land. Its nearest extant relatives are two
species, P. taiwanensis from Taiwan and
P. hwangshanensis from mainland China
(considered by most Chinese botanists
to be synonymous), indicating a migration route from the SW to the islands. Due to its tolerance
of salt-laden ocean winds P. luchuensis can grow close to the shore avoiding competition from
other trees; however it also occurs as an emergent tree at higher altitudes to 700 m in mixed
angiosperm-coniferous forest, together with the podocarps Nageia nagi and Podocarpus macro-
phyllus. On Okinawa, the largest island, coastal and lowland pine forests were over-exploited and
then replaced by plantations after the Second World War; remaining original stands are now
protected from logging.
 mainland asia and japan 273

Map MAJ-157. Pinus amamiana [n=6, m=3, h=2, 1914–2008]

Pinus amamiana has a very limited distribution and

occurs only on two islands to the S of Kyushu, Yakush-
ima and Tanegashima. It is closely related to some
other E Asian pines in subsection Strobi and exhibits
adaptive traits in its seed cones and seeds that indicate
dispersal of the seeds by birds. Its nearest relatives are
uncertain as this species has as yet not been included
in DNA analyses so we do not know whether it is closer
to Japanese or to mainland Asian species. Pinus ama-
miana occurs mostly in rocky terrain at low elevations
between 100–300 m in sparsely vegetated habitat. In
total, not more than 3000 mature trees appear to exist, 2000 of them on Yakushima. Due to exploi-
tation in the past and slow growth the population is declining; an accidentally introduced species
of nematode also seems to cause problems with increased seedling mortality. This species is listed
as Endangered (EN) on the IUCN Red List.

Map MAJ-158. Pinus parviflora var. parvi-

flora (black) [n=59, m=50, h=12, 1862–2003];
P. parviflora var. pentaphylla (red) [n=33,
m=31, h=8, 1892–1988]

Two varieties are distinguished within

this species, Pinus parviflora var. parvi-
flora and var. pentaphylla. Pinus parvi-
flora is widespread in Japan, ocurring on
the four main islands and on Okushiri
Island off the W end of Hokkaido. Out-
side Japan it is only known to occur
on the mountain Jiri-san (Chiri-san) in
South Korea and on Ullung-do (Dagelet
Island), an island in the Sea of Japan
ca. 125 km E of and belonging to South
Korea. The two varieties have overlapping distributions but var. pentaphylla is more prevalent in
thee N of Honshu and in Hokkaido and is absent in Shikoku, Kyushu and in Korea. This species
is also often planted as an ornamental tree; such records are excluded from the map. Pinus parvi-
flora occurs at altitudes from just above sea level to about 2500 m, with an optimum between 1000
and 1500 m a.s.l. in montane forests. This pine often retreats to steep, rocky slopes and outcrops
where the dense montane forest is less well developed or does not reach, thereby avoiding com-
petition from broad-leaved angiosperm trees. At high, subalpine locations it becomes a dwarfed
tree but unlike the more northerly distributed species P. pumila (map MAJ-144 on p. 264 it does
not develop a layering habit.

Picea

The genus Picea has six species in Japan and five of these are endemic to Japan. The sixth species,
P. jezoensis, is more widespread in the Far East of N Asia, but one of its subspecies, P. jezoensis
ssp. hondoensis, only occurs in Honshu. In two other endemic spruces of Japan, varieties are
recognized. This is somewhat exceptional for a genus with many widespread species and may
indicate a higher rate of speciation in the genus having occurred in the central part of Honshu,
where most of the endemic spruces occur.
274 mainland asia and japan

Map MAJ-159. Picea jezoensis ssp. jezoensis var.

jezoensis (black) [n=44, m=41, h=11, 1893–2003];
P. jezoensis ssp. jezoensis var. komarovii (red)
[n=6, m=6, h=4, 1918–1986]; P. jezoensis ssp. hon-
doensis (yellow) [n=39, m=39, h=8, 1862–2003]

Picea jezoensis is a variable species with a

‘maritime’ distribution around the Sea of
Japan and in a few areas further NE, the
most notable being an occurrence in cen-
tral Kamchatka, which Russian botanists
tend to classify as a separate species. Her-
barium collections from the coast of the Sea
of Okhotsk and from Sakhalin are sparse
(or were not at our disposal) so the map is
incomplete for these areas. Reports of occurrences further inland are doubtful if not backed by
verified herbarium specimens. Picea jezoensis appears to be present on two of the the three Kuril
Islands nearest to Japan (Hokkaido), Kunashi and Iturup (herbarium specimen mapped); fur-
ther out towards Kamchatka the islands have a subarctic climate and mostly treeless vegetation.
Milder climatic conditions in the past could have provided a ‘stepping stone’ migration route.
Picea jezoensis ssp. jezoensis var. jezoensis is the northern form of this species and forms extensive
forests in Hokkaido ( Japan), Sakhalin, Primoriye (few data available) and Kamchatka. The forest
in the broad, low altitude valley of the Kamchatka River is a taiga relict with P. jezoensis as the
dominant conifer, locally mixed with Larix gmelinii var. gmelinii (map MAJ-146 on p. 266 Pinus
pumila in the sub-canopy, and Betula. In Jilin Province, China and adjacent North Korea occurs
Picea jezoensis ssp. jezoensis var. komarovii. Presumably var. jezoensis occurs also in North Korea,
but no data are available; it is present in at least two localities in South Korea (two of E. H. Wil-
son’s collections at K in the Conifer Database from N. Hankyo Prov. could not be located). Picea
jezoensis ssp. hondoensis occurs in Honshu (Hondo) in Japan, most abundantly in the central part
of the Japanese Alps, the largest area is in Nagano, Shizuoka and Yamanashi Prefectures and a
second area to the NE in Gunma and Tochigi Prefectures. Two outlying locations are to the N in
Iwashiro Prefecture (Komaga-dake) and a larger area to the S in Nara Prefecture. This subspecies
is montane and occurs between 1100 m and 2700 m a.s.l. where it is scattered in mixed deciduous
angiosperm-coniferous forest with Abies, Tsuga and Pinus (all evergreen as is Picea) and Larix
(deciduous like the angiosperm trees).

Map MAJ-160. Picea glehnii [n=21, m=21, h=10,

1861–1983]

Picea glehnii occurs in Hokkaido and Honshu

(Japan) and in Sakhalin and the Kuril Islands
(Russia). Its widest distribution is in Hokkaido,
occurring in many localities across the island,
where it is mostly a constituent of mixed
conifer forest with Abies sachalinensis, Picea
jezoensis and as a lesser component Taxus
cuspidata at higher elevations to ca. 1600 m. It
becomes a more scattered tree below ca. 800
m down to near sea level in deciduous angio-
sperm forest. In Sachalin, from where the spe-
cies was first described, it appears to be rare
 mainland asia and japan 275

and found only in the S of the island. In Honshu a very small population is known since 1960 to
occur in Iwate-ken in the NE near the Pacific coast in mountains at 1000–1100 m a.s.l. (Schmidt-
Vogt, 1987, f. 40 [map]) but it is not shown on our map as no herbarium specimens were available.
A small population also occurs on Kunajiri Island in the Kuril Islands chain (Russia), the large
island of this chain nearest to Hokkaido. It is possibly present on some of the smaller islands to
the S of Kunajiri Island but no specimen data are available.

Map MAJ-161. Picea maximowiczii var. maxi-

mowiczii (black) [n=10, m=9, h=12, 1864–1965];
P. maximowiczii var. senanensis (red triangle)
[n=2, m=2, h=3, 1965–1965]

This is one of the rarest species of Picea in

Japan with a restricted range mostly in Nagano
Prefecture, central Honshu. The locality of the
type collection made by Tschonoski (Chôno-
suka Sugawa) in 1864 was either given as “Prov.
Senano” (= Shinano) (GH) or “Fudzi-yama”
(= Tokura-yama?) (P) or not at all (other
herbaria). Note that “Fudzi-yama” is not the
famous Fuji-yama (Fuji-san) which is much
further S in Yamanashi Prefecture, where
this species does not occur. Yet the location
given here (the most W point on the map)
remains tentative. Most localities are in the Kanto-sanchi and Yatsuga-dake Mountains some
distance NE of this putative type locality. Two varieties are recognized, with var. senanensis the
rarest and located farthest E but apparently partly sympatric with var. maximowiczii. This spe-
cies occurs in the mountains between 1100 m and 2000 m a.s.l. in small, scattered groups, usually
associated with Juniperus rigida, sometimes with Pinus densiflora and various broad-leaved trees,
mostly in very open, grassy terrain. The variety senanensis has been found with Picea alcoquiana
and P. koyamae. Exploitation and habitat degradation have caused serious decline of this slow
growing species and both varieties are listed as Endangered (EN) on the IUCN Red List.

Map MAJ-162. Picea koyamae [n=6, m=6, h=5,

1911–2005]

Picea koyamae has a very limited distribution

in the Akaishi and Yatsuga-dake Mountains,
Nagano Prefecture, Honshu. Most of the six
herbarium collections found for the Atlas
are from Mt. Yatsugadake itself, one on the
S-slope, the others unspecified. This spe-
cies occurs in very small stands, usually of
less than 20 mature trees, and is most often
mixed with Larix kaempferi, Picea alcoqui-
ana var. acicularis, P. maximowiczii in a few
localities, and various broad-leaved trees
and shrubs. It is limited to N-facing slopes
at 1500–2000 m altitude. Due to the fact that
probably no more than a few hundred trees exist in largely unprotected and (formerly) exploited
mountain forests this species is listed as Critically Endangered (CR) on the IUCN Red List.
276 mainland asia and japan

Map MAJ-163. Picea torano [n=32,

m=32, h=9, 1889–2004]

Picea torano (syn. P. polita) is

widespread in Honshu, Shikoku
and Kyushu; it is the Japanese
spruce that reaches furthest S
and is consequently adapted to
a warmer climate than the other
species in Japan. The northern-
most occurrence is in Fukushima-
shi in N Honshu at 37° 45’ N, for
which a herbarium collection was
found to give a dot on the map.
According to a similar map in
Kurata (1964, vol. 1: 26 as P. pol-
ita) this species is also common
in Shikoku and Kyushu but from
each island we could find only a single herbarium collection. It appears that this species does not
occur in the mountains on the Sea of Japan side (NW side) of Honshu; in winter cold air and snow
come from the NW across this sea, while the Pacific side of Honshu generally stays warmer. The
altitude at which this spruce occurs ranges from 400 m to 1600 m so it stays at lower elevation
than other species of Picea in the central mountains of Honshu. It also rarely forms pure stands
and is more commonly found in mixed conifer forest with Abies homolepis, Larix kaempferi, Tsuga
sieboldii (mainly in the S of its range) or mixed with deciduous angiosperm trees. Past exploita-
tion has reduced the number of trees in many areas and P. torano is listed as Vulnerable (VU) on
the IUCN Red List.

Map MAJ-164. Picea alcoquiana [n=15, m=13, h=8, 1861–1974]

Picea alcoquiana has a limited distribution in central

Honshu, mainly in the Kanto and Chubu Mountains.
Three varieties are recognized but not separated on the
map for lack of herbarium specimens with unambigu-
ous identifications. The populations of this spruce are
small and scattered and occur usually in mixed conif-
erous forest with Picea jezoensis ssp. hondoensis as the
most common of the spruces, Tsuga diversifolia and
Larix kaempferi, both also common, Pinus parviflora
and Abies veitchii in some areas, A. mariesii usually at
higher elevations, and broad-leaved trees, e.g. Betula
ermanii, B. grossa, Sorbus commixta, Quercus mongol-
ica var. grosseserrata, Alnus hirsuta var. sibirica, and Prunus maximowiczii. The altitude range is
considerable, from 700 m to 2180 m and the composition of the forest changes from angiosperm
dominated to conifer dominated with increasing altitude. The two rarest varieties, P. alcoquiana
var. acicularis and var. reflexa, are both listed as Endangered (EN) on the IUCN Red List.

Pseudotsuga

Only a single species of Pseudotsuga occurs in Japan. A map of the genus, which has just four
species in total, is presented in the chapter North America (map. NA-44 on p. 59).
 mainland asia and japan 277

Map MAJ-165. Pseudotsuga japonica

[n=10, m=8, h=9, 1895–1980]

Pseudotsuga japonica is a rare spe-

cies with a limited distribution,
mostly confined to the Kii Peninsula
in SW Honshu but with a few very
small subpopulations in the E part of
Shikoku. It occurs scattered in mixed
forests at altitudes between 500 m
and 1000 m on steep slopes or in
ravines, often in rocky places where
the forest is less well developed. It
is a relatively small, broad-crowned
tree not able to compete with taller
trees in the forest. Abies firma and Tsuga sieboldii are the most common conifers in these forests;
several species of Quercus dominate the angiosperm element. The total population is estimated to
be fewer than 2000 mature trees and the species is in decline, resulting in a listing as Endangered
(EN) on the IUCN Red List.

Pseudotsuga japonica (grey-green trees) in mixed woodland with Abies firma, Pinus densiflora, Sciadopitys
verticillata and Tsuga sieboldii on the Kii Peinisula, S Honshu. Photo credit and © Dendrological Atlas Team
(Zsolt Debreczy & István Rácz).

Larix

The genus Larix is represented by two taxa in Japan, Larix kaempferi and Larix gmelinii var. japon-
ica. The first species is endemic to Honshu, Japan but Larix gmelinii is a widespread and variable
species in NE Asia, of which the var. japonica is not endemic to Japan and occurs in Hokkaido
(Japan), Sakhalin and the Kuril Islands (both Russia). We present it therefore with the other variet-
ies of L. gmelinii in the section Mainland Asia Far East of this chapter (map MAJ-146 on p. 266).
278 mainland asia and japan

Map MAJ-166. Larix kaempferi [n=38,

m=37, h=6, 1862–2005]

Larix kaempferi is widespread and

common in the mountains of cen-
tral Honshu, but its natural extent
of occurrence (EOO) is actually
rather limited, as it is present in
just seven prefectures, ranging
from 35° 10’ N to 36° 52’ N and from
137° 30’ E to 139° 45’ E according
to herbarium specimen records in
the Conifer Database. Larix kae-
mpferi is a species of mesic sites,
occurring from the hills to high
in the mountains (500 m to 2300
m a.s.l.), on the south face of Fuji
san it reaches 2900 m. It is com-
monly found in association with
other conifers, e.g. Pinus densi-
flora, Picea jezoensis subsp. hon-
doensis, Tsuga diversifolia, Abies
homolepis at lower elevations, and
A. veitchii at higher elevations, but it is clearly a sub-climax species. Generally growing on fertile
soils of volcanic origin and capable of rapid growth, L. kaempferi has turned out to be a desirable
tree in forestry and it has been planted elsewhere in Japan as well as in many other countries,
especially in W Europe.

Tsuga

The genus Tsuga has nine species, of which five are native in Asia and four are North American. It
is one of several genera with an E Asian/North American distribution. Its global distribution divides
into three main areas, W North America, E North America and E Asia (Farjon, 1990: 150). The genus
is absent in the boreal taiga of Alaska-Canada, Scandinavia and Russia-Siberia; its northern limits
(60° 10’ in Alaska, 48° 40’ in New Brunswick, Canada, 40° 13’ N in Japan) are all in maritime envi-
ronments and the most continental occurrence is of T. canadensis in Minnesota, USA (map NA-52
on p. 64). Further disjunction of the range of this genus separates the two endemic species of Japan
from T. chinensis in Taiwan and this island population is again separated from scattered occurrences
of the same species in mainland China and a larger area with three species in S-Central China and
the Himalaya. In W North America the populations of two species are divided into a Pacific coast
range and a Rocky Mountain range (maps NA-50 and NA-51 on pp. 62–63). This discontinuous dis-
tribution of Tsuga, a forest forming conifer, is apparently a remnant of a much wider range of the
genus in the geological past. In Eurasia, fossils are known from the Eocene through the Pliocene
and during the younger epoch Tsuga still occurred in Western and Central Europe, S Russia, W and
E Siberia and Japan (Florin, 1963). During the Eocene it was also present in the maritime provinces
of the Russian Far East; from the Oligocene through the Pliocene it existed in W North America,
including areas where it is now absent (Axelrod, 1976). The present disjunctions are probably the
result of range contractions from N latitudes in the interior parts of the Northern Hemisphere con-
tinents and of extinctions, particularly in Europe, caused by gradual cooling since the Early Miocene
and more abrupt and severe periods of cooling in the Pleistocene.
 mainland asia and japan 279

Phylogenetic analyses of all species have resulted in two or three clades (Havill et al., 2008; Les-
lie et al., 2012) which separate three North American species from an Asian clade which however
contains T. caroliniana from the Appalachian Mountains in E North America. Estimated divergence
time for the split between Nothotsuga and Tsuga falls in the Late Cretaceous to earlier Paleogene
and the divergence time for the two clades dividing the genus Tsuga is estimated to be later Paleo-
gene/Neogene. Diversification of species in the Asian clade occurred during the Neogene, i.e. in the
Early Miocene or later. This would imply that there were still connections between Asia and North
America involving Tsuga during the Miocene, presumably via the Bering Land Bridge, and that
there was no early divergence separating the genus on each continent. Tsuga caroliniana may have
migrated from Asia to North America via the BLB (Havill et al., 2008).

Map MAJ-167. The distribution of the genus Tsuga is disjunct, with three main areas, two in North America
and one in E Asia.

Map MAJ-168. Tsuga sieboldii [n=35,

m=35, h=15, 1842–2006]

Tsuga sieboldii occurs in central

Honshu, Shikoku and Kyushu,
on the island of Yakushima S of
Kyushu, and outside Japan only
on Ullung-do (Dagelet Island) in
the Sea of Japan, ca. 125 km E of
South Korea. In Honshu, it is rare
or absent in Nagano Prefecture
and adjacent parts of the Japanese
Alps and no herbarium records
were found from the most W part
of Honshu. The distribution of
this species therefore appears to
be discontinuous and the occur-
rence on Yakushima and espe-
cially on Ullung-do are probably
due to chance dispersal of seeds.
Tsuga sieboldii grows in hills and mountains at altitudes between 400 m and 1500 m a.s.l. (from
500 m to 950 m on Shikoku). It is usually associated with conifers such as Abies firma, Pseudotsuga
japonica, Chamaecyparis obtusa, Cryptomeria japonica, Pinus densiflora, P. parviflora, and Sciado-
pitys verticillata, and with various angiosperm trees in mixed forests. Exploitation of this conifer
has been quite intense but it is still considered common in its main areas of occurrence.
280 mainland asia and japan

Map MAJ-169. Tsuga diversifolia [n=56,

m=53, h=8, 1863–2002]

Tsuga diversifolia has a more northerly

distribution than T. sieboldii (map MAJ-
168 on p. 279 as it is rare in Shikoku and
Kyushu and in SW Honshu only occurs
on the Kii Peninsula, while it occurs in N
Honshu where T. sieboldii is absent. Its
main distribution is in central Honshu,
particularly in Nagano, Tochigi, Gunma
and Yamanashi Prefectures. These pre-
fectures encompass the Japanese Alps,
the complex mountain system of the
central part of the main island of Japan.
In the conifer forests of these moun-
tains Tsuga diversifolia is often the most
common species, but several other conifers can occur with it. Tsuga diversifolia is shade tolerant and
could eventually prevail over many other species but is checked by episodic disturbances (or man-
made disturbances such as logging) that under natural conditions allow other species to regenerate
by opening up the canopy. This species can become dominant especially in very wet climate, where
year-round precipitation creates ‘mossy forest’ with a dense undergrowth of Rhododendron sp. or
even blanket bogs in which it becomes shrubby. The altitude range of T. diversifolia is 700–2000 m,
rarely higher and lowest in the N of Honshu.

Abies

The genus Abies is present in Japan with five species, four of which are endemic. Of the fifth spe-
cies, A. sachalinensis, var. mayriana is also endemic, occurring only in Hokkaido. The genus is
widespread in Japan and occurs on all four main islands and on several of the smaller islands as
well. Several species are abundant and major constituents of coniferous forests or indeed domi-
nant, especially at higher altitudes in the mountains.

Map MAJ-170. Abies firma [n=72, m=70, h=12,

1842–2005]

Abies firma is the most common and

ubiquitous fir in the warmer parts of
Japan, occurring in Honshu, Shikoku
and Kyushu as well as on Yakushima to
the S of Kyushu and on the islands of
Tsushima in the Korean Strait. In Hon-
shu it is distributed across much of the
island as far N as Miyagi Prefecture at
ca. 38° 30’ N, but this species is rare in
the mountains facing the Sea of Japan,
which receive much snow during winter
storms. Abies firma occurs on hills and in
mountains at elevations between 50 and
1200 m a.s.l. (commonly 300–1000 m) and
forms extensive conifer forests with Tsuga sieboldii on higher slopes but is mostly found in mixed
conifer-angiosperm forests together with many other species of trees.
 mainland asia and japan 281

Map MAJ-171. Abies mariesii [n=65, m=61, h=9,

1878–2005]

Abies mariesii is an endemic fir of Honshu, the

largest of the Japanese islands. This species has
a more northerly distribution than Abies firma
(map MAJ-170 on p. 280) occurring from central
Honshu, where the ranges of the two species
overlap, to Aomori Prefecture in the northern-
most part of the island. This division is simi-
lar to the distributions of Tsuga diversifolia
(map MAJ-169 on p. 280) and T. sieboldii (map
MAJ-168 on p. 279) and reflects climatic changes
between SW and NE Honshu from warmer to colder winters. Abies mariesii is a species of the
high mountain sides and ridges in the upper montane and subalpine zones, occurring commonly
between 1000 m and 2800 m a.s.l. (as low as 750 m in N Honshu). It occurs above A. firma where
the ranges of the two species coincide and at the highest altitudes it can form pure forest stands
up to the tree line. In most localities it is part of a mixed high altitude coniferous forest with
Abies veitchii, Tsuga diversifolia, Picea jezoensis var. hondoensis and scattered Betula and Acer as
the most common angiosperm trees.

Map MAJ-172. Abies veitchii var. veitchii (black) [n=40,

m=38, h=7, 1910–2005]; A. veitchii var. sikokiana (red)
[n=7, m=7, h=4, 1965–1984]

Abies veitchii is another endemic fir of Japan, in

which species two varieties are recognized. Of
these, A. veitchii var. veitchii is the most common
and widespread, with its main distribution in the
Japanese Alps in the centre of Honshu and a few
outlying localities to the NE. Another disjunct
occurrence of this variety is on the Kii Peninsula
in an area called Tenkawa-mura (Nara Prefec-
ture). Abies veitchii var. sikokiana occurs in three
locations in Shikoku, on Mt. Ishizuchi (Tengu-dake), Mt. Tsurugi and on nearby Mt. Sasagamine.
This species is a montane to subalpine fir occurring from 1200 m to 2800 m a.s.l. as a co-dominant
tree in mixed coniferous forests, usually together with A. mariesii, A. homolepis, Tsuga diversifolia,
Picea jezoensis, Larix kaempferi, Thuja standishii, Pinus parviflora and other trees including angio-
sperms such as Acer and Betula.

Map MAJ-173. Abies homolepis [n=44, m=42, h=9,

1842–2005]

The distribution of Abies homolepis is very similar

to that of A. veitchii, with its main area of occur-
rence in central Honshu in the Japanese Alps,
a smaller area on the Kii Peninsula (where it
appears to be more widespread than A. veitchii)
and Mt. Ishizuchi and Mt. Tsurugi in Shikoku.
On these two mountains and on many others in
central Honshu both firs are found together. Its
altitude range is somewhat lower than that of
A. veitchii with 1100 m to 1800 m in Shikoku and
282 mainland asia and japan

700 m to 2000 m in Honshu, but there is considerable overlap in the zone where a mixed conif-
erous forest is predominant. At lower altitude another fir, A. firma becomes the dominant spe-
cies, and here a mixed deciduous-evergreen forest with angiosperms such as Fagus, Acer and
­Carpinus ­providing the deciduous element while the firs and other conifers are evergreen. In
autumn, when the leaves of the angiosperm trees turn yellow and red, the dark green conifers
provide a striking contrast.

Map MAJ-174. Abies sachalinensis

[n=21, m=20, h=10, 1862–2003]

Abies sachalinensis occurs pri-

marily in Hokkaido (Japan) and
in Sakhalin (Russia) and also on
a few smaller islands, of which
we have herbarium records on
Moneron Island E of the S end
of Sakhalin and on Iturup Island
(Yeterofu) in the S Kuril Islands.
There is also an enigmatic occur-
rence on the Pacific coast of
Kamchatka. Several varieties have
been recognized in this variable
species, but the herbarium data
are too limited to allow them to be mapped with the exception of var. gracilis in Kamchataka
(red circle). We present a map of the species here. The occurrence in Sakhalin is more wide-
spread than shown on the map, but few herbarium collections have come to our attention. In the
Kuril Islands it is probably present on the islands nearest to Hokkaido, but again there was just
one herbarium collection available to us. The occurrence in Kamchatka, originally described as a
new species Abies gracilis, is controversial (Farjon, 2010a). The firs occur in a completely isolated
‘grove’ mixed with Betula on a coastal lowland site near an abandoned native village; there is no
other conifer forest along this coast. Is this a natural population or a case of introduction to cre-
ate a ‘sacred’ grove of evergreen trees in a treeless landscape by people who traveled from the S
along the Kuril Islands? Is it a species, a variety or not even distinct from A. sachalinensis? This
very disjunct grove of firs far to the N of any nearest occurrence of the genus Abies in E Asia is
suspect, but only a thorough study combining archeology, genetics and ecology could provide an
answer.

Abies sachalinensis in central Hokkaido at ca. 700 m, with sporadic Picea jezoensis. Photo credit and
© Dendrological Atlas Team (Zsolt Debreczy & István Rácz).
 mainland asia and japan 283

Podocarpaceae

The family Podocarpaceae reaches its northern limit in Japan (for the distribution of the family
see map GTC-16 on p. 21). Only three species are native in Japan, one in the genus Nageia and
two in Podocarpus. All three have distributions that extend beyond Japan to Taiwan and/or main-
land China. Of these, only Podocarpus fasciculus has its main distribution in Japan, the other two
species, Nageia nagi and Podocarpus macrophyllus, have much wider distributions in China and
Taiwan and are treated in that section of this chapter.

Map MAJ-175. Podocarpus
fasciculus [n=26, m=25,
h=7, 1853–2008]

Podocarpus fasciculus is
closely related and simi-
lar to P. macrophyllus and
has been confused with
that species. It occurs
in the Ryukyu Islands
between Kyushu and Tai-
wan and on some islands
S of Kyushu (among
which is Yakushima) and
on a peninsula on the W
coast of of Kyushu, as well
as in Taiwan. It is possibly
present on other islands
near Kyushu but herbar-
ium data have not been
available, in part due to
identification issues. In
the Ryukyu Islands this
species occurs in low
canopy rainforest in localities sheltered from ocean winds and remains a small tree; in Taiwan it
grows taller and occurs in mixed evergreen rainforest with conifers and angiosperms at altitudes
between 1500 m and 2500 m. Podocarpus fasciculus is listed as Vulnerable (VU) on the IUCN
Red List.

Sciadopityaceae

Japan uniquely is the country with an endemic family of conifers, Sciadopityaceae, which con-
tains only a single species, Sciadopitys verticillata. This taxon is sister to a clade that contains the
family Cupressaceae, now including the formerly recognized family Taxodiaceae. Fossils ascribed
to Sciadopityaceae date back to the Late Jurassic of Europe and well preserved cones of Scia-
dopitys from the Middle to Upper Cretaceous are known from Japan (Stockey et al. in Farjon,
2005a). Leaves and cone fragments that belong to Sciadopitys have been described from Eocene
to Pliocene sediments in Europe and demonstrate that this peculiar conifer was once widespread
in Eurasia and experienced a severe range contraction during the Pleistocene. There were once
several distinct species and earlier in the Mesozoic Era perhaps distinct genera.
284 mainland asia and japan

Map MAJ-176. Sciadopitys verticillata [n=44, m=34,

h=16, 1775–1983]

Sciadopitys verticillata is distributed in Hon-

shu, Shikoku and Kyushu. It is most abun-
dant in Honshu on the Kii Peninsula in Nara
and Wakayama Prefectures and in Shikoku.
In Kyushu it is only known from Miyazaki
Prefecture. Other natural occurrences are in
the central mountains of Honshu in Gifu and
Nagano Prefectures. The mountain Koya-san
in Wakayama Prefecture has one of the larg-
est stands of S. verticillata with several ancient
trees. This unique conifer has been planted
widely near shrines and temples and it is not
always possible from herbarium collection
data to distinguish those occurrences from specimens collected from trees growing in natural
habitat. Its northernmost natural locality is probably Nozawa-machi in Fukushima Prefecture at
37° 35’ N which is indicated on the map. Another interesting locality is Nii-jima Island in the Izu
Seven Islands group S of the Izu Peninsula, also indicated on the map. Here S. verticillata grows as
stunted, flat-topped trees on tuffstone cliffs almost directly above the Pacific Ocean. This conifer
is more commonly found in rocky, cool and moist ravines and valleys in mountainous areas, at
altitudes between 200 m and 1700 m a.s.l. where it is associated with several other species e.g.
Chamaecyparis obtusa, C. pisifera, Tsuga sieboldii, Abies firma and Pinus parviflora and various
angiosperm trees.

Cupressaceae

The family Cupressaceae is represented in Japan with the genera Cryptomeria, Thujopsis (both
monospecific and endemic to Japan), Thuja (1 species, endemic), Chamaecyparis (2 species, 1
endemic, 1 shared with Taiwan) and Juniperus (5 species, 2 endemic). Of these species, two are
much more widespread outside Japan, with Juniperus chinensis mapped under China and Taiwan
in this chapter (map MAJ-98 on p. 237) and J. communis treated in the chapter Europe and the
Mediterranean (maps EM-31-32 on pp. 160–161).

Cryptomeria

Map MAJ-177. Cryptomeria japonica [n=52, m=24,

h=12, 17??–1997]

The natural distribution of Cryptomeria

japonica, the national tree of Japan and that
country’s most important timber tree by far,
is extremely difficult to establish. Natural for-
ests that include this species are now very
rare (Tomaru et al., 1994)and those forests in
which it still occurs have been greatly altered.
This potentially very large conifer has been
both logged and (re-)planted for many cen-
turies. Outside Yakushima Island all existing
large and old trees have been planted. Ernest
Wilson (Wilson, 1916) after his travels in
 mainland asia and japan 285

Japan in 1914 concluded that no natural forests of/with this conifer existed apart from the forest
on Yakushima, which he visited. It is perhaps not as serious as that; there are forests elsewhere
in which both plantations and spontaneously growing trees of this species occur, quite likely
these are sometimes on sites which had ‘old growth’ i.e. uncut natural stands of C. japonica in
the past. The species was also exported to China long ago; it is not native there despite claims to
the contrary and the history of its planting in China is actually quite well documented in Chinese
literature. Areas where Cryptomeria japonica may still be indigenous are, apart from Yakushima,
the Kii Peninsula in S-central Honshu and several localities on the coast of the Sea of Japan, e.g.
in Akita Prefecture and on Sado Island. This species in natural forest is an emergent, long living
tree that resembles Fitzroya cupressoides and Sequoiadendron giganteum in its ecology and long-
term life cycle. This cycle is artificially shortened in managed forests.

Cryptomeria japonica below Kuromi-dake in Yakushima, with scattered Chamaecyparis obtusa and Tsuga
sieboldii. Photo credit and © Dendrological Atlas Team (Zsolt Debreczy & István Rácz).

Thujopsis

Map MAJ-178. Thujopsis dolabrata

var. dolabrata (black) [n=76, m=71,
h=12, 1775–2006]; T. dolabrata var.
hondae (red) [n=27, m=23, h=8,
1899–2005]

This endemic species is wide-

spread and occurs on all four
main islands of Japan, from the
Oshima Peninsula in Hokkaido
to Kyushu. Of the two varieties,
T. dolabrata var. dolabrata is the
most widespread and it is only
in N Honshu that var. hondae
appears and finally replaces the
nominate variety in Aomori Pre-
fecture and S Hokkaido. Thujopsis
dolabrata var. dolabrata is also
found on a few smaller islands,
especially Sadoga-shima in the Sea of Japan. Small populations occur on Kozu-shima and Nii-
jima in the Izu Seven Islands group S of the Izu Peninsula in thee Pacific Ocean. In the far SW of
286 mainland asia and japan

Japan this species becomes rare and it is apparently absent on Yakushima, an island with a varied
conifer flora of at least eight species. Thujopsis dolabrata is a relatively shade-tolerant conifer and
occurs at altitudes from 400 m in the N to 2100 m in the S in mixed conifer-angiosperm forest or
mixed conifer forest, in the furthest northern parts of its range sometimes in pure stands, spread-
ing through layering.

Thuja

The genus Thuja has five species, two in North America and three in E Asia. The two species in
North America are separated in E and W distributions; in Asia the three species also have disjunct
ranges, with one species in Japan, one in Korea and adjacent Jilin Province of China and one in
a very small area in Chongqin Municipality, central China. This distribution is an example of the
well known Asia-North America disjunct distribution of trees, involving taxa or their immedi-
ate ancestors which had a much wider distribution across the Northern Hemisphere up to the
Pliocene or Early Pleistocene. Fossils from the High Arctic in Ellesmere Island, Canada and from
Spitsbergen indicate a wide distribution of Thuja at high latitudes well within the Arctic Circle
during the earlier Cenozoic (Palaeocene-Eocene). Younger fossils are known from Sikhote Alin in
the Russian Far East and from Japan. Phylogenetically Thuja is most closely related to Thujopsis,
now a monospecific genus endemic to Japan (map MAJ-178 on p. 285) and of which there is as
yet no fossil evidence that it occurred elsewhere in the geological past. The present distribution
of Thuja and its separation into five species, two of them with very limited occurrence, is most
likely the result of range contraction and vicariance.

Map MAJ-179. The distribution of the genus Thuja.

Map MAJ-180. Thuja standishii [n=67, m=59, h=14,

18??–2005]

Thuja standishii is widespread and common in

the mountains of central Honshu but rare in
the SW of the island and in Shikoku. No her-
barium collections are known from Shikoku.
It occurs northward in Honshu as far as the
mountains of Akita Prefecture at ca. 40° 10’
N. Thuja standishii is also present on Okino-
shima in the Sea of Japan. Its altitudinal
range is (250–)500–2000(–2500) m a.s.l. and
so it occurs, usually as a minor component,
in both mixed angiosperm-conifer forest and
in mixed conifer forest. Often it is confined to
moist rocky precipices facing N, sites less suit-
able for larger trees.
 mainland asia and japan 287

Chamaecyparis

The genus Chamaecyparis has five species with a disjunct distribution in North America with
two species and in Asia (Japan and Taiwan) with three species. The American species have also
highly disjunct ranges, with one species in a limited area on the West Coast (N California and
SW Oregon) and the other species widespread on the Atlantic Coastal Plain. In Asia the genus is
absent on the mainland but widespread in Japan and Taiwan. Japan and Taiwan both have one
endemic species and share a third, albeit with a distinction at varietal rank between the two ter-
ritories. This is another example of the well-known Asia-North America disjunction in plants and
especially trees. The fossil record gives evidence of a much wider distribution in North America
from the Late Cretaceous to the Miocene, with High Arctic presence in the warmest epoch, the
Eocene. No unambiguous fossils are known from Europe (Stockey et al. in Farjon, 2005a), but fos-
sil taxa that may be closely related, e.g. Mesocyparis, were apparently widespread in the Northern
Hemisphere from the Late Cretaceous to the Eocene or later. The phylogeny of Chamaecyparis
appears to indicate closer relationship of the E North American C. thyoides with the Japanese spe-
cies C. pisifera than with its W North American congener. This clade diverged from the remainder
of the species before these again separated, suggesting vicariance events taking place in different
periods of geological time separating more widespread ancestral taxa (Wang et al., 2003).

Map MAJ-181. The distribution of the genus Chamaecyparis.

Map MAJ-182. Chamaecyparis pisifera [n=50,

m=46, h=9, 1800–2005]

Chamaecyparis pisifera occurs in Hon-

shu from Aomori-shi in the N to Hyogo
Prefecture in the W. It is also present
in a few localities in Kyushu but has
not been found in Shikoku and in the
most western part of Honshu. The near-
est to Shikoku is a population on the
island Awajishima. An outlying popula-
tion occurs on Mikura-jima, one of the
Izu Seven Islands group in the Pacific
Ocean. Chamaecyparis pisifera occurs
at altitudes between 280 m and 2600 m in wet places with high rainfall or water close to the
surface. Its ability to layer in these circumstances causes it to spread and form clonal groves or
stands, but it is scattered and can be absent in large areas within its extent of occurrence (EOO).
Due to its wide altitude range it occurs in various forest zones from lowland deciduous forest
dominated by angiosperms to high montane evergreen coniferous forest, but in both it tends to
occupy sites less favourable to tree growth.
288 mainland asia and japan

Map MAJ-183. Chamaecyparis obtusa var. obtusa (left) [n=80, m=75, h=9, 1800–2006]; Map MAJ-184. Chamae-
cyparis obtusa var. formosana (right) [n=31, m=26, h=12, 1900–2003]

These two maps show the disjunct distribution of Chamaecyparis obtusa and the two populations
are commonly recognized as distinct varieties. Chamaecyparis obtusa var. obtusa is widespread
in central and W Honshu and in Shikoku and occurs more scattered in Kyushu. It is also present
on several smaller islands, e.g. Tsushima in the Korea Strait, Shimo-koshiki-shima W of Kyushu
and Yakushima S of Kyushu. The northern limit of this Japanese variety is at ca. 37° 10’ in Fuku-
shima Prefecture. It occurs in mixed forests at altitudes between 80 m and 2500 m which means
in lowland angiosperm forest as well as high montane conifer forest and transitional forest types.
In some of these, especially on dryer sites, it can grow to a large, emergent tree. Chamaecyparis
obtusa var. formosana is endemic to Taiwan, where it mainly occurs in the central mountains. It
can become a large tree to 50 m tall and is locally common in mixed conifer forest with C. formo-
sensis, Taiwania cryptomerioides, Cunninghamia konishhi and Calocedrus formosensis and several
angiosperm trees and shrubs at altitudes between 1800 m and 3000 m. High levels of precipita-
tion to 4000 mm and more per annum and daily fog make these conifer forests among the most
lush and well developed in E Asia. Extensive logging of these forests as well as their conversion
to plantations of Cryptomeria japonica and other fast growing conifers have reduced the area of
occupancy. This variety is listed as Vulnerable (VU) on the IUCN Red List.

Juniperus

Five species of Juniperus are native to Japan and two of these are endemic, while three occur
also in mainland Asia (or even beyond). Of the latter, one is mapped here as it has a nearly
endemic subspecies; the other two occur mostly outside Japan and, as indicated above on p. 284
are treated elsewhere.

Map MAJ-185. Juniperus rigida ssp. conferta

(red) [n=31, m=27, h=10, 1800–1997]; J. rigida ssp.
rigida (black) [n=74, m=62, h=11, 1800–2006]

Juniperus rigida ssp. conferta is wide-

spread in Japan and also occurs in Sakha-
lin, Russian Far East. This subspecies is a
decumbent shrub which is strictly littoral
and occurs on sandy ocean shores, form-
ing dense, spreading mats of shrubby
 mainland asia and japan 289

v­ egetation covering old beaches and dunes. Usually the vegetation is very open and has a pioneer
character, but this juniper will sometimes occur (persist) in Pinus woodland near the shore. It has
been found in Sakhalin by E. H. Wilson in August 1914 (Wilson 7365 at K), but without indication
of locality, so its place on the map is tentatively put at Yuzhno-Sakhalinsk which he would have
visited sailing from Hokkaido. This subspecies occurs from there along the shores of Japan to as
far as Amami-shima in the Ryukyu Islands (another Wilson collection: Wilson 8199 of March 1917
at K). Other islands from which it was collected are Tanega-shima S of Kyushu and O-shima just
E of the Izu Peninsula. Juniperus rigida ssp. rigida also occurs in Japan but is widespread on the
mainland of E Asia, from S Primorye in the Russian Far East through Korea and into NE China
as far W as the Helan Shan in the border area between Ninxia and Nei Mongol (Inner Mongolia).
This subspecies is an erect shrub or small tree and can be found on the coast but then usually
on higher slopes of hills or mountains; more often it occurs in the interior in mountains to 2200
m a.s.l. in various types of more or less open vegetation and on a great diversity of soils, from
moorland or sand dunes to rocky mountain slopes and limestone cliffs.

Map MAJ-186. Juniperus taxifolia

[n=24, m=22, h=9, 1827–1977]

Juniperus taxifolia has an

extremely disjunct distribution
as it occurs in the Ryukyu Islands
(Nansei-shoto), in the Ogasawara
Group of the Bonin Islands, and
in two localities on the S coast of
Honshu. This distribution forms
a triangle with sides of ca. 1500 ×
1500 × 800 km between the fur-
thest points of occurrence, cross-
ing open stretches of the Pacific
Ocean. The two localities in S Honshu are on the coasts of the Izu and Bozo Peninsulas. Juniperus
taxifolia forms a prostrate or decumbent shrub close to the seashore above the tide mark and
also occurs on rocks or rocky slopes from sea level to the highest parts of some of the islands. It
is also present further inland, where it can be a small, erect tree, in tall grassland and thickets
on deeper, though usually rocky or gravely soils. Its peculiar ‘maritime’ distribution on oceanic
islands separated by wide spaces of ocean suggests dispersal by birds that visit (or visited) these
islands, but no specifics are known about these dispersers.

Juniperus taxifolia on Chichi-jima in the Bonin Islands,

Japan. The natural occurrence of this and two other species
of Juniperus i.e. J. bermudiana and J. brevifolia on oceanic
islands far from major land masses is evidence for seed
dispersal by birds across long distances. Dependence on
migratory birds seems to be the sine qua non for the colo-
nisation of such islands by conifers; these chance events
have left most oceanic islands uninhabited. Photo credit
Rafael Govaerts.
290 mainland asia and japan

Map MAJ-187. Juniperus procumbens [n=7,

m=6, h=3, 1800–1917]

Juniperus procumbens is another Japa-

nese juniper that is remotely dispersed
on small islands. Some of these are near
Ryukyu (Iki-shima, Ojiki-shima and Shi-
kano-shima) and other islands are in the
distant Ryukyu Islands chain (Okinawa in
the Okinawa Group and Ishigaki in the
Sakishima Group). This species, in Japan
by some botanists regarded as a variety
of J. chinensis and perhaps because of
this concept sometimes confused with
shrubby forms of that species occurring
in Honshu, is apparently rare and only
occurs on small islands, similar to J. taxifo-
lia but not as widely dispersed. The ecol-
ogy of J. procumbens is poorly known but
most specimens were collected not on the
sea shore but inland, creeping over rocks,
sometimes at considerable altitude. Its
true extent of occurrence may likewise
be incompletely known and it could be
found on other islands lying between the
farthest points known at present.

Cephalotaxaceae

In Japan, a single species of Cephalotaxus occurs with two varieties. The majority of species in
this genus occur in China and the genus is mapped and discussed under China and Taiwan in
this chapter.

Map MAJ-188. Cephalotaxus var. harringtonii

(black) [n=65, m=61, h=9, 1839–2007]; C. har-
ringtonii var. nana (red) [n=51, m=50, h=5,
1926–2006]; C. harringtonii var. wilsoniana
(purple) [n=18, m=18, h=6, 1912–2002]

Cephalotaxus harringtonii occurs in

Japan, South Korea and Taiwan. In
Japan, two varieties are present, C. har-
ringtonii var. harringtonii and var. nana.
In South Korea only var. harringtonii has
been found to grow in natural habitat. In
Taiwan, C. harringtonii var. wilsoniana is
the only taxon in the genus occurring in
the wild; it is sometimes treated as a dis-
tinct species C. wilsoniana. [The species
is often alternatively spelled as C. har-
ringtonia.] This species extends farthest
 mainland asia and japan 291

NE of all species in the genus, reaching NW Hokkaido at ca. 44° 20’ N and 141° 45’ E; most species
occur in warm-temperate to subtropical regions of SE Asia. In Japan, var. nana occurs farthest
N and is also the most common variety on the NW side of Honshu, where in winter most of
the snowfall occurs and stays for longer than elsewhere (‘snow belt’). The nominate variety har-
ringtonii is more common on the Pacific side and in the SW of Honshu; in Kyushu, Tsushima and
South Korea it is the only variety present. This separation is not absolute and both shrub (var.
nana) and tree (var. harringtonii) may sometimes occur on the same mountain, but then often at
different altitudes or exposures. Cephalotaxus harringtonii occurs from near sea level on the coast
to ca. 1900 m in the mountains of Honshu. The shrub form can grow in thickets with Taxus cuspi-
data var. nana and Torreya nucifera, or form extensive layering clones under canopy of deciduous
angiosperm trees, while var. harringtonii occurs as a sub-canopy tree in mixed forest or conifer
forest. In Taiwan var. wilsoniana is a small tree occurring in mixed forest at 1800–2700 m a.s.l.

Taxaceae

The family Taxaceae is represented in Japan by two species in different genera: Taxus cuspidata
and Torreya nucifera. Both also occur outside Japan, but Torreya nucifera is a near endemic as it
is widespread in Japan and only occurs outside that country on a few South Korean islands.

Taxus

Map MAJ-189. Taxus cuspidata [n=78, m=77, h=13, 1888–2010]

Taxus cuspidata occurs in lands around the Sea of Japan; its main range is in Japan from where
it extends NE on to the Kuril Islands and N to the S part of Sakhalin. On the Asian mainland it
is present in Primorye (Russian Far East), especially in the hills and mountains near Vladivostok,
in the S part of Jilin and Liaoning Provinces of China, and in Korea where it reaches as far S as
Cheju-do (Quelpart Island). Taxus cuspidata also occurs in S Heilongjiang Province and has been
reported from Shaanxi Province (both China) in Flora of China vol. 4 (Fu et al., 1999) but from the
latter province, far to the SW, no correctly identified herbarium specimens were traced and it may
be introduced there. In Korea it is rare and occurs in a few isolated mountain localities, usually as
very old trees with little or no regeneration evident. Two varieties are recognized, T. cuspidata var.
cuspidata and var. nana, the latter a low shrub, in Honshu often occurring on rocky slopes high
above the Sea of Japan but also in the interior on mountain ridges; they are not separated on the
map because herbarium specimens are often not identified at varietal rank. The species occurs at
altitudes between 100 m and 1600 m in deciduous to semi-evergreen angiosperm forest in the S
and in coniferous forest in the N of its range and is a minor component in the lower canopy.
292 mainland asia and japan

Torreya

Map MAJ-190. Torreya nucifera

[n=108, m=104, h=16, 1712–2005]

Torreya nucifera is widespread

and common in Japan from
Akita Prefecture in N Honshu
to Kyushu; it is also present on
Cheju-do (Quelpart Island) and
on Wando Island of South Korea.
It is reported from the island of
Yakushima but no herbarium
collection was found to place it
on the map. In N Honshu it is
confined on the W-side of the
island but southward from ca. 38°
N it spreads to both coasts and
becomes common in both angiosperm-dominated and mixed angiosperm-conifer forest. The alti-
tude ranges from near sea level to ca. 1100 m. Old-growth forest with ancient trees of T. nucifera
in large numbers occurs in Korea’s Bija-Rim Forest (Halla-san National Park, Cheju-do) with trees
estimated to be 500–800 years old. A more or less decumbent form of this conifer develops in the
mountains of Honshu facing the Sea of Japan, where snowstorms from the W can pile up 8–10 m
of snow on the lee side of ridges. Cephalotaxus harringtonii var. nana and Taxus cuspidata var.
nana often occur with Torreya nucifera in this exposed habitat; the habit they have developed is
an adaptation to deep snow cover.
MALESIA

Geography

Malesia in this Atlas is the region in tropical SE Asia covered by Flora Malesiana (up to 1988 when
conifers were treated), with one addition, the island of Bougainville, which is geographically part
of the Solomon Islands but politically belongs to Papua New Guinea. This region includes the
following countries: Brunei, East Timor, Indonesia, Malaysia, Papua New Guinea, Philippines and
Singapore. The total land area is 3,021,630 km² and is made up of a vast archipelago extending on
either side of the Equator from the Malay Peninsula in the west to Bougainville in the east and
from islands in the Luzon Strait between Taiwan and the Philippines in the north to the island of
Timor in the south. The only area in the region connected to the mainland of Asia is Peninsular
Malaysia which forms the southernmost part of the Malay Peninsula. The largest islands are New
Guinea, Borneo, Sumatera, Jawa, Sulawesi, Luzon and Mindanao. Islands in a second size class are
situated in the Philippines, the Moluccas and the Lesser Sunda Islands. The seas around the islands
of the archipelago are significant in the interpretation of the distribution of conifers, due to the
fact that large parts of the Sunda Shelf fell dry during glacial maxima of the Pleistocene, connect-
ing the Malay Peninsula with Sumatra, Java and Borneo, while deep sections of ocean kept other
islands and archipelagos isolated. Similarly, New Guinea became connected with Australia across
the Sahul Shelf and while sea straits remained between the Philippines and Taiwan and mainland
China, they became narrower. These colder periods lasted longer than the interglacials with high
sea levels as at present and therefore these ‘bridges’ with adjacent floral regions may have influ-
enced distribution patterns recognized today. Van Steenis (1950) found that the ‘bridges’ despite
their relatively recent history of connections still form principal floristic ‘demarcation knots’ setting
the flora of Malesia apart from adjacent regions. The deep sea sections and straits that maintained
divisions between islands and archipelagos have caused internal divisions of biodiversity, princi-
pally separating West Malesia from East Malesia along different ‘lines’. The best known of these is
Wallace’s Line which separates the Philippines, Borneo and the Greater Sunda Islands + Bali from
Sulawesi, the Lesser Sunda Islands (– Bali), Maluku and New Guinea. There are mountain ranges
and systems on all the major islands and the Malay Peninsula and many of these are volcanic in
origin. Mt. Kinabalu in Sabah, Borneo (4094 m) is the highest summit between Yunnan in China
and New Guinea (Mt. Jaya, 5030 m). Extensive lowlands are situated in E Sumatra, S Borneo and
S New Guinea, in many places with swamps and slow rivers. The climate is monsoonal throughout,
but most markedly so in E Jawa and the Lesser Sunda Islands, while the highest mountains in the
Philippines, Mt. Kinabalu and New Guinea are wet throughout the year.

Conifers in Malesia

Families Genera Species Species + infra Countries with Area of occupancy Endemic taxa
conifers in km²
6/8 13/70 80/615 85/794 7/7 26,850/3,021,630 63
Numbers after ‘/’ are global or continent totals.
294 malesia

Conifers occur naturally throughout Malesia, with species present on the Malay Peninsula and all
the major islands and many of the smaller islands as well.

Map MA-1. Distribution of conifer species in Malesia.

Most species occur in mountains and consequently a greater density of dots on the map is found
where the land is mountainous, while the major lowland areas mentioned above have very few
conifer species, with large areas apparently without any conifers. The map has to be interpreted
with some caution, as there is certainly a collection bias against some areas. For instance, the
density of dots in the western (Indonesian) part of New Guinea is less than in the eastern (Papua
New Guinean) part, not because there are necessarily fewer conifers but because that half of the
island has not been as intensively surveyed botanically as PNG. Similarly, the Indonesian part
of Borneo (Kalimantan) may be as diverse in conifers as Sarawak to the N of it, especially in the
borderlands, but fewer herbarium collections have been made there. A similar bias may exist
for parts of the Philippines, but here a long history of deforestation could have caused the low
density of dots in e.g. Mindanao, Panay and Leyte. The Lesser Sunda Islands E of Flores, including
Sumba to the south, are also poor in conifers or do not have them at all. This is genuine, caused by
climate; these islands are closest to Australia from which a high pressure cell during July–August
intensifies the dry season from June to September. All Malesian conifers are adapted to humidity
and not to dryness. The lack of conifers on these islands is matched in the tropical zone of the
Northern Territory in Australia (map AU-4 on p. 345) just 400 km across the Timor and Arafura
Seas. The two principal ‘bridge’ areas for conifers are the Malay Peninsula, with species extend-
ing into mainland SE Asia and S China, and Bougainville/Solomon Islands connecting with the
SW Pacific islands. The other ‘bridge’ across Torres Strait to Australia has resulted in only one
taxon occurring on both sides, Sundacarpus amarus. The land on either side of the strait has no
conifers for hundreds of km, so the gap at least at present is wider than it would seem. Malesia
is therefore relatively isolated and being so extensive and topographically diverse naturally has a
high level of endemism, 74% of conifer taxa occur only here. Distributions vary from across the
region and beyond to a single mountain. In the latter category, Mt. Kinabalu stands out with four
species endemic to that mountain. The long and high ‘backbone’ of mountains in New Guinea
has many conifer species not present elsewhere or only on the nearest islands on either end.
Conifer diversity in Malesia is substantial with 80 species but not high compared to Australasia
with 119 species. While isolation may have played its part in speciation, none of the islands are as
 malesia 295

isolated as New Caledonia and New Zealand, both with 100% endemism for conifers. Because of
this greater connectivity, local endemics on such large islands as Borneo and New Guinea exist
alongside species with a wider distribution. Wallace’s Line running midway though the archipel-
ago seems to have little significance for conifer distributions in Malesia as distribution boundaries
rarely coincide with it while many species cross it. Due to these facts, it is not really informative
to divide this chapter in geographical sections and we will instead present the species by family
and genus, but grouping them more or less by their pattern of distribution. The endemic species
of Mt. Kinabalu, all in Podocarpaceae, are discussed together and the following species, with
wider distributions outside Malesia, are mapped and discussed in other chapters: Pinus kesiya,
Taxus wallichiana (maps MAJ-135 on p. 257 and MAJ-33 on p. 195), Araucaria cunninghamii and
Podocarpus insularis (maps AU-10 on p. 350 and AU-125 on p. 426

�‒�
�‒�
�
�‒�
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Map MA-2. Map of Malesia showing numbers of conifer genera per 0.5 degree cell.

�‒�
�‒�
�‒��
��‒��
��‒��

Map MA-3. Map of Malesia showing numbers of conifer species per 0.5 degree cell.
296 malesia

The two maps MA-2 and MA-3 are showing taxonomic diversity distributions at generic and spe-
cific ranks respectively. The largest number of 0.5 degree cells with high diversity is found in New
Guinea, particularly at the generic level. This is due to a nearly consistent high level of diversity
along the main mountain chain of the island and in the Arfak Mountains in the Bird’s Head
Peninsula. Smaller concentrations are in N Borneo and, scoring slightly less, in Peninsular Malay-
sia, N Sumatera and Luzon in the Philippines. Grid cells with fairly high generic diversity (6–7
genera) occur in several places outside New Guinea, e.g. Peninsular Malesia, Borneo, Sulawesi and
the Philippines. High species diversity is much less widely distributed than high generic diversity;
outside New Guinea there are just two other cells in Peninsular Malesia and in the Philippines
scoring 12–15 species. Sabah stands out for species diversity in the whole of Malesia with two cells
scoring 16–19 species, mostly on account of Mt. Kinabalu. While generic diversity is somewhat
more evenly distributed, speciation has really taken off on the high mountains of Sabah, in par-
ticular Mt. Kinabalu. Given its young geological age, this diversification is likely to have occurred
comparatively recently. Diversity at both taxonomic ranks is associated with mountanous areas;
the majority of conifer taxa occur in the mountains of Malesia.

Map MA-4. Rarity values per 0.5 degree cell. Cell score is the total of the rarity calculations for each species
in the cell. The rarity calculation is 1 divided by the total of cells occupied by a species. Thus, a species that
occurs in one cell scores 1 while a species that occurs in 10 cells scores only 0.1.

Map MA-4 gives the distribution of narrow endemism, species with limited distribution and where
these are concentrated. Unlike in some other regions, e.g. Australasia and South America, in Male-
sia endemism and diversity occur largely in the same areas: New Guinea, Borneo, N Sumatera and
Peninsular Malesia. Exceptions are Luzon in the Philippines, which scored high in generic diver-
sity, and Sulawesi, which appears to have no endemic conifer species at all. Narrowly endemic
species in Malesia are nearly all montane, with a few exceptions e.g. Nageia maxima (map MA-57
on p. 324) and possibly Podocarpus costalis (map MA-14 on p. 302 which is somewhat wider dis-
tributed. In Malesia, as will be discussed in the relevant cases, several species are only known
from the type locality and perhaps one or two other. This does not necessarily mean that they
are so restricted, our knowledge may be deficient and these species may occur elsewhere. If this
were the case ‘artificial’ endemism will colour the map. We can only map what we know and only
further field surveys looking out for these species can resolve that problem.
 malesia 297

Podocarpaceae

The family Podocarpaceae is by far the most diverse family in Malesia, with seven genera (7/13) and
65 species (65/80). The three genera Podocarpus (31 species), Dacrydium (15 species) and Dacrycar-
pus (9 species) make up 85% of species in the family. The remainder are five species in Falcatifo-
lium, three species in Nageia, one species in Retrophyllum and one species in Sundacarpus.

Map MA-5. The distribution of species of Podocarpaceae (black) and of species belonging to other families
(red) in Malesia.

When the map of all species (map MA-1 on p. 294) is divided into those belonging to Podocar-
paceae and all other conifer species, some interesting patterns emerge. Podocarpaceae, as can
be expected from the statistics given above, occupy the same general area as all conifers, i.e.
they are present in the Malay Peninsula and all major islands (first and second size class) of
the region. However, they are the only conifers naturally present in Jawa and the Lesser Sunda
Islands. Conversely, Podocarpaceae appear to be rare in Kalimantan, the Indonesian part of
Borneo. This region is under collected, but given the predominance of the Podocarpaceae over-
all, the bias should be on the side of this family, not against it, if species of this family were
more common there.

Podocarpus

The largest genus of the family Podocarpaceae (31 species in Malesia) is widely distributed
in the region. The map of the genus is very similar to the map of the family. In some areas,
such as the Malay Peninsula and N Sumatera, Podocarpus appears to be less common than
other genera such as Dacrydium; in other regions, e.g. New Guinea, species of Podocarpus
seem to predominate. The overall impression is that of full occupation of the archipelago by
the genus.
298 malesia

Map MA-6. The distribution of the genus Podocarpus in Malesia.

Map MA-7. Podocar-
pus neriifolius var. neri-
ifolius [n=325, m=276,
h=27, 1821–2009]

Podocarpus neriifolius
has the widest distri-
bution of all species
in the genus, extend-
ing from NE India
and S China to Fiji
in the SW Pacific. In
Fiji, both var. neriifo-
lius and var. degenerii
occur; the latter vari-
ety has been treated in the chapter Australasia (map AU-124, p. 426). In China, P. neriifolius is most
common in Hainan, N Guangzhou, Guangxi and N Hunan, but it occurs in other provinces
as far as Sichuan and SE Xizang (Tibet). Its most western occurrence is in the hills near Dar-
jeeling, India and from there it is distributed eastward all through Indochina and down the
Malay Peninsula into Malesia as far as the Bismarck Archipelago and the Louisiades Archi-
pelago. Further eastwards, it occurs in the Solomon Islands, Vanuatu and Fiji. The distance
from Darjeeling to Fiji is around 10,000 km and therefore the range of Podocarpus neriifolius
is in the same category as that of Pinus sylvestris and only surpassed by Juniperus communis,
both of which are essentially boreal species. Podocarpus neriifolius has its main distribution
in the tropics, but extends in mainland Asia into warm temperate forests. This remarkable
range, as well as the common occurrence of P. neriifolius in evergreen primary broad-leaved
forests, in China from 100–1500 m and in Malesia up to 2900 m a.s.l., is evidence of a highly
successful tropical conifer.
 malesia 299

Podocarpus neriifolius var. neriifolius. A sapling tree in the Nan Ling Mts. of Guangdong Province, China
(left). Photo credit Aljos Farjon. A tall tree left standing in deforested hills, New Guinea and its ‘fruits’ (seed
+ receptacle). Photo credit Tim Utteridge.

Map MA-8. Podocarpus rubens [n=35, m=35, h=6, 1878–2000]

Podocarpus rubens has a wide but scattered distribution in Malesia. It is a species of mon-
tane evergreen rainforest, attaining 45 m in optimal growing sites, but at high altitude in New
Guinea usually stunted as other trees in ‘mossy forest’. On most of the large islands of Malesia
only a few collections have been made that were identified as this species. This rarity could be
in part due to identification problems; the leaves of juvenile trees resemble smallish leaves of
mature trees of P. neriifolius and when tree size was not stated with the collected herbarium
specimen, it may be taken to belong to the much more common species, which shares the
same habitat.
300 malesia

Map MA-9. Podocarpus polys-

tachyus [n=110, m=99, h=17,
1822–1998]

Podocarpus polystachyus is
common in Peninsular Malay-
sia, on islands between Sumat-
era and Borneo, along the W
and N coast of Borneo, on
Palawan Island and in Luzon;
elsewhere it has been found
more sporadically, the furthest
east in the western lowlands
of New Guinea near Lobo. It
is curiously very rare on the
E coast of Sumatera. Podocarpus polystachyus is also commonly planted, usually as an amenity
tree, and these occurences have been excluded from the map as much as possible, depending on
information given with the herbarium specimens. It is most often found in the coastal lowlands
on sandy soil, often directly inland from the zone of mangroves on old dunes or beaches, and
in coastal ‘kerangas’ forest, now increasingly being replaced by oil palm plantations. Estimated
decline of this habitat has caused P. polystachyus to be listed as Vulnerable (VU) on the IUCN
Red List. Further inland, it is found on eroded limestone escarpments or plateaus. Its altitude
range is from near sea level to 500 m, but on Palawan it has been found at 1000 m.

Map MA-10. Podocarpus
pilgeri [n=99, m=98, h=21,
1895–2008]

Podocarpus pilgeri has

a very wide distribution
extending from S China to
the Solomon Islands. It is
common in New Guinea,
the Philippines and on
the limestone karst moun-
tains of N Viet Nam and
adjacent parts of Yunnan
and Guangxi in China. It
is rare in Borneo (Sarawak) and uncommon in Sulawesi. It seems to have been collected in the
Moluccas only once, but this may be due to under-collecting. Podocarpus pilgeri is conspicu-
ously absent from the Greater and Lesser Sunda Islands and from the Malay Peninsula. It is a
small-leaved species which occurs in montane forests (often ‘mossy forest’) and in low scrub
on exposed mountain ridges and summits. Its altitudinal range is (700–)1200 to 3000(–3300) m
a.s.l. While it can be a large tree, it is often smaller or a shrub. Low mossy Xanthomyrtus-Podo-
carpus forest dominates the crests and summit areas of Mt. Hunstein (Papua New Guinea) and
Mt. Nettoti (Papua, Arfak Mts.) where Podocarpus pilgeri is a decumbent shrub. At its highest
point (3300 m in New Guinea) it grows on the edge of low, open ‘heath forest’ and subalpine
grassland on peaty soil.
 malesia 301

Map MA-11. Podocarpus rumphii [n=91, m=82, h=16, 1891–1993]

Podocarpus rumphii is another widely distributed species in Malesia. It is most common in New
Guinea and the Moluccas to the NE tip of Sulawesi and occurs more scattered elsewhere, including
on some of the smaller islands. This species has been reported from Hainan Island in China, but
no herbarium collection could be found to substantiate its presence there. Specimens are also lack-
ing from Sumatera and the Bismarck Archipelago. Sterile foliage specimens of this species can be
confused with large leaves from sapling trees of the widespread and common species P. neriifolius,
although the leaves of P. rumphii tend to be wider than these. [Herbarium labels should always men-
tion the size and estimated age of the tree sampled and whether it grew in shade or sun.] Podocarpus
rumphii occurs in lowland to lower montane rainforest, usually below 500 m but occasionally up to
1600 m a.s.l. It grows on a variety of soils, including lowland clays, limestone and acidic rock types
and attains 45 m under good conditions, successfully competing with angiosperms (including Dip-
terocarpaceae) probably because of its large leaves, especially on sapling and pole trees in shaded
conditions. These trees occur in low densities with slow regeneration from solitary seedlings.

Map MA-12. Podocarpus bractea-

tus [n=21, m=20, h=4, 1888–1973]

Podocarpus bracteatus is the

only species in the genus with
a distribution limited to the
Sunda Islands. It occurs in N
Sumatera, in Jawa and on the
island of Flores. The Sunda
Islands have a significantly
lower diversity of conifer gen-
era and species than the other
islands and Peninsular Malay-
sia (maps MA-2 and MA-3 on p. 295). Podocarpus bracteatus is a scattered canopy tree of ever-
green montane tropical rainforest, most commonly found on forested volcanoes of W Jawa (Jawa
Barat) at altitudes between 1000 m and 2600 m but occasionally at lower elevation.
302 malesia

Map MA-13. Podocarpus
glaucus [n=23, m=23, h=6,
1906–1984]

Podocarpus glaucus is
mainly found E of Wal-
lace’s Line but is also
known from Mt. Halcon in
Mindoro, Philippines. Most
herbarium collections are
from New Guinea, the Bis-
marck Archipelago and the
Solomon Islands; two are
from Sulawesi and Seram.
Podocarpus glaucus is
most commonly a decumbent or erect shrub occurring in stunted ‘mossy forest’ on mountain ridges,
either of acidic rock or karst limestone. It can also occur in forest on the edge of high altitude peat
swamps, where it occasionally forms a small tree. Such habitats are not rare W of Wallace’s Line and
yet apart from the Mt. Halcon population it has not been found there. Presumably, this species had
an eastern origin and the occurrence in the Philippines is the result of a dispersal event.

Map MA-14. Podocarpus costalis [n=12, m=10, h=9, 1907–1998]

Podocarpus costalis is a shrubby tree occurring on Lanyu (Orchid

Island) E of the southern end of Taiwan and on several small islands
in the Luzon Strait (Batan Islands, Babuyan Group) belonging to the
Philippines. It also occurs in a few places on the E coast of Luzon and
on Palasan Island in Quezon Province. In Taiwan and in Luzon this
attractive podocarp is planted and also used for bonsai; its natural
occurrence on the N coast of Luzon is in doubt because of this; in Tai-
wan occurrences on the mainland are invariably planted or at least
introduced. Its natural habitat is limestone bluffs and sea stacks, where
it occurs from near sea level to 300 m in lowland evergreen scrub. Its
specific habitat and few localities where it occurs in the wild make it a
rare species. Indiscriminate collecting of live plants to be transplanted
to gardens have caused decline of populations and Podocarpus costalis
is now listed as Endangered (EN) on the IUCN Red List.

Map MA-15. Podocarpus teysmannii

[n=25, m=22, h=8, 1855–1990]

Podocarpus teysmannii occurs in

Peninsular Malaysia, Sumatera
(including Bangka and Lingga
Islands) and Borneo (Brunei,
Sabah). It seems peculiarly absent
from Sarawak, perhaps this is an
artefact of collecting efforts. It is
a sometimes tall but usually small
tree of both primary and second-
ary lowland rain forest, often on
sandy or rocky ridges with poor
 malesia 303

soil, or in ‘kerangas’ forest. In the latter it grows with Dryobalanops rappa (Dipterocarpaceae),
Agathis sp., Casuarina sp. and small-leaved shrubs on often wet sand in coastal areas; this is a
vegetation type now increasingly converted to oil palm plantations. On Gunung Benum (Pahang,
Peninsular Malaysia) P. teysmannii was collected at 1140 m and on the Mesilau River in Mt. Kina-
balu National Park, Sabah it occurs at 1350 m a.s.l. so not all populations are in the lowlands.

Map MA-16. Podocarpus deflexus (red) [n=12, m=12, h=7, 1905–1973]; Podocarpus levis (black) [n=16, m=16,
h=5, 1872–1979]

Podocarpus deflexus is only known from two locations, Gunung Tahan in Pahang, Peninsular
Malaysia and the Gunung Leuser Reserve, Aceh, N Sumatera. It is a conspicuous species with very
long and narrow, deflexed leaves so if it occurs elsewhere, it is unlikely to have been missed. On
the other hand, while the Gunung Tahan population is represented by more than 10 herbarium
collections in the Conifer Database, dating from 1905 to 1973, the Sumatran locality is known
from just one at the Leiden herbarium (L) made in 1972. Both populations are within protected
areas, but habitat degradation is an issue in the Gunung Leuser Reserve. This species is listed as
Endangered (EN) on the IUCN Red List. Podocarpus levis is mainly distributed E of Wallace’s Line
in Sulawesi, the Moluccas and W New Guinea, but with one locality in Borneo. Little is know
of its habitat other than ‘evergreen primary rainforest’ from near sea level to 1650 m according
to herbarium collection data, and that the Bornean population grows on limestone. Its taxo-
nomic status is similarly uncertain (Farjon, 2010a) and if distinct, it may have been confused with
P. neriifolius in the field.

Map MA-17. Podocarpus ridleyi [n=20, m=19, h=6,

1894–1978]

Podocarpus ridleyi is restricted to Peninsular Malaysia

(including Penang Island) where it is most common
on mountain summits such as Gunung Ledang (Mt.
Ophir), Gunung Blumit, Gunung Jerai, Kedah Peak and
Gunung Panti. Below the summit areas it can be found
on ridges down to around 500 m a.s.l. but it is absent
on densely forested mountain slopes unless on impov-
erished soils derived from sandstone or granite (Mt.
Ophir) where the forest is thinner and more stunted.
These localized habitats were formerly islands in low-
land rainforest, now they are increasingly islands of
natural vegetation in a sea of oil palm and rubber tree
plantations. Even so, these island habitats are not safe
unless effectively protected and Podocarpus ridleyi is
now listed as Vulnerable (VU) on the IUCN Red List.
304 malesia

Map MA-18. Podocarpus globulus [n=10, m=10, h=7,

1960–1979]

Podocarpus globulus has a limited distribution in

N Borneo (Sabah and N Sarawak) where it occurs
in isolated populations on a few mountains. Her-
barium collections were made on Bukit Ampuan,
Bukit Tawai and Gunung Silam in Sabah and in the
Maligan Range in Lawas District, Sarawak. The most
common habitat is ‘mossy forest’ on steep moun-
tain ridges, but this species has also been found in
rainforest on ultrabasic igneous rock. This species is
listed as Endangered (EN) on the IUCN Red List.

Map MA-19. Podocarpus borneensis [n=7, m=7, h=7,

1875–1996]

Podocarpus borneensis is only known with certainty

from four localities in Borneo: Bukit Tawai in Sabah,
Bario and “Buta Laga” near Kapit, both in Sarawak,
and Karimata Island off Kalimantan Barat. In Flora
Malesiana 10 (3) (De Laubenfels, 1988) a map is given
on p. 403 with seven localities, but no information
on what sources this is based and the herbaria A, K,
KEP, L, MO and US only have seven collections in all,
four from D. J. de Laubenfels (P698, P700–702), from
the four localities here mentioned. Specifically, no
collections were found from the Merurong Plateau
in Sarawak, where it was said to be common. This, too is a species from ‘mossy forest’ on mountain
ridges, but is also reported from ‘kerangas’ forest on sandy soils.

Map MA-20. Podocarpus laubenfelsii [n=15, m=15, h=7,

1956–2005]

Podocarpus laubenfelsii occurs on Mt. Kinabalu and

on Mt. Trus Madi in Sabah, in Lawas District and
Pulong Tau National Park, Sarawak and on Gunung
Palimasan in Kalimantan Timur. A herbarium col-
lection for the locality “Kutei” in Kalimantan Timur,
mentioned and mapped in Flora Malesiana 10 (3)
(De Laubenfels, 1988) has not been traced. This spe-
cies grows scattered in ‘keranga’ forest with Agathis
borneensis, Nageia wallichiana, Sundacarpus ama-
rus, Dacrydium gracile, and Falcatifolium falciforme,
often on nutrient-poor and/or water-logged, acidic
soils. The species is also scattered in primary rain-
forest and ‘mossy forest’, the latter on exposed mountain ridges. Deforestation is destroying the
habitat of this species especially in Sabah outside protected areas and it is listed as Endangered
(EN) on the IUCN Red List.
 malesia 305

Map MA-21. Podocarpus confertus [n=14, m=14, h=7,

1957–1979]

Podocarpus confertus is another rare species of

Podocarpus from Borneo. Podocarpus confer-
tus has been collected on Bukit Ampuan, on
Gunung Silam and in Kuala Penyu in Sabah
and on limestone hills in the Bau District of
Sarawak. The Kuala Penyu population near the
coast in W Sabah is at 20 m altitude, while the
more often collected Bukit Ampuan popula-
tion is on mountain ridges at around 1200 m.
Gunung Silam is a small mountain near the E
coast of Sabah in the Bay of Lahad Datu. The
habitat is usually described as ‘kerangas’ forest,
but on the mountain ridges it should be ‘mossy
forest’ where many conifers retreat from competition with large-leaved angiosperms lower down
the slopes. This species is listed as Endangered (EN) on the IUCN Red List.

Map MA-22. Podocarpus micropedunculatus [n=20,

m=20, h=5, 1957–1996]

Podocarpus micropedunculatus is restricted to

N Borneo, where it is mostly found in Brunei,
but also in the Marudi Forest Reserve of N Sar-
awak and in the Kimanis and Mandahan For-
est Reserves on the South China Sea coast of
Sabah. An isolated occurrence is in the Tawau
Hills in SE Sabah. This species occurs on wet
sandy or peaty soils in ‘kerangas’ forest or along
forest edges, where it forms shrubby thickets.
Under canopy of Agathis sp. it often grows to a
small tree. It shares with two Australian species
a capacity of rhizomatous propagation, but it is
not known if here on the coastal lowlands of N
Borneo this is equally an adaptation to fire.

Map MA-23. Podocarpus brassii [n=68, m=68, h=5, 1913–2000] Podocarpus brassii at high altitude is a dense
shrub with small, thick leaves; the large single seed and receptacle are ready to be taken by a bird while
others around it were aborted. Photo credit Timothy Waters.
306 malesia

Podocarpus brassii has two varieties, var. brassii and var. humilis, which are primarily distinguished
by features of their pollen cones. As dioecious plants, half of the living individuals will be difficult
to identify to variety under any circumstances, but with much of the herbarium material sterile it
becomes an almost impossible task to give an accurate map of the two varieties. They are therefore
ignored in this map. Podocarpus brassii is a species of the central mountain chain of New Guinea,
occurring from the Daniau Panai Lakes (‘Wissel Lakes’) in the west to Goe in the Owen Stanley
Range in the east. It occurs at high altitudes, from 2500 m to 4000 m a.s.l., in ‘mossy forest’ or high
montane forest, in subalpine scrub and marginally in alpine tussock grassland, variously forming a
small tree or a spreading, low shrub. The forest line is often situated around 3650 m but tree forms
are not always limited to altitudes below this and edaphic factors also play a role, with wet and
boggy sites inhibiting tree growth and instead favouring dense mats of the low, shrubby form.

Map MA-24. Podocarpus ledermannii [n=21, m=21,

h=7, 1912–1989]

Podocarpus ledermannii is widespread in New

Guinea and the Bismarck Archipelago, but it is
relatively rare, only 21 collections were found
in the institutional herbaria holding specimens
from New Guinea. This species occurs from near
sea level, e.g. in river deltas at Purari and Sepik,
to 2300 m in tropical montane evergreen rainforest and is said to be sometimes locally common in
Castanopsis forests. While there may be an artefact of collecting, it is notable that on the map it is as
rare in Papua New Guinea, where collecting has been more intense, as in West New Guinea (Papua).

Map MA-25. Podocarpus crassigemma [n=32, m=32,

h=5, 1954–1989]

Podocarpus crassigemma is distributed in

Papua New Guinea and possibly also in W New
Guinea (Papua) where Flora Malesiana 10 (3)
(De Laubenfels, 1988) gives two dots on the
map in fig. 58 on p. 413 but for which no her-
barium collections were found in A, K or L. There is a collection (K, L) made in 1967 in Papua on
or near the border, shown on our map. Martin Sands (K) found it in 1975 on Mt. Angil in New
Ireland and a New Guinea Forestry (NGF) collection (K) was made in 1973 in the Sulu Range in
New Britain; neither are present in the Flora Malesiana map. Some of these discrepancies are
due to incorrect determinations of herbarium collections. Most collections are from the Eastern
Highlands (Bismarck Range) around Goroka where Podocarpus crassigemma is a common large
canopy tree in montane rainforest at 2400–2650 m with other Podocarpus spp. and angiosperms
of the families Cunoniaceae, Fagaceae, Lauraceae, Myrtaceae, and others. At higher altitudes up
to 3400 m it occurs in ‘mossy forest’ dominated by Nothofagus and may extend to the subalpine
zone with tree ferns and tussock grasses.

Map MA-26. Podocarpus pseudobracteatus [n=26,

m=26, h=8, 1933–1989]

Podocarpus pseudobracteatus has a similar

distribution to that of P. crassigemma but
does not occur in the Bismarck Archipelago.
It is most common in the Southern and East-
ern Highlands of Papua New Guinea where it
 malesia 307

occurs in montane ‘mossy forest’ dominated by Castanopsis and Nothofagus, at altitudes between
2200 and 2850 m. At higher altitudes it enters dwarfed forest and scrub bordering on subalpine
peat lands and tussock grassland. The two known localities in the Indonesian (western) half of
New Guinea are in the Maoke Range near Lake Habbema and on the slopes of Peak Trikora.
Whether these are indeed the only two localities in that great mountain range is doubtful, but if
not Podocarpus pseudobracteatus is undoubtedly much less common in W New Guinea than in
E New Guinea.

Map MA-27. Podocarpus archboldii [n=24, m=24,

h=8, 1939–1985]

Podocarpus archboldii occurs across the island

of New Guinea in the mountains. Outside the
highlands of Papua New Guinea it is only known
from Bernhard Camp on the Idenburg River in
the northern foothills of the Pegunungan Maoke
Range and from the Kebar Valley in the Arfak
Mountains of the Birds Head Peninsula. We have once more the difficult distinction between lim-
ited collecting compared to Papua New Guinea and genuine rarity to consider. Several specimens
in CANB from PNG under this name belong to P. crassigemma instead, but have not been renamed.
Subtracting these and mapping those actually belonging to P. archboldii still gives many more col-
lections in PNG and so we must conclude that it is probably more common there than in Papua.
This species is a canopy tree of montane evergreen rain forest occurring from 1500 m to 2600 m,
with a few collections made lower down to 720 m a.s.l. It is therefore geographically sympatric with
P. crassigemma but occurs mostly at lower elevations, with only a few hundred meters of overlap.
Podocarpus archboldii is listed as Vulnerable (VU) on the IUCN Red List.

Doubtful Species and/or Distributions

Six species of the genus Podocarpus in Malesia are either of uncertain distinction or insufficiently
known to map with confidence. These species are (with indications where they might belong taxo-
nomically) Podocarpus macrocarpus (= P. rumphii ?), P. palawanensis (= P. polystachyus ?), P. ramosii
(= P. pilgeri ?), P. atjehensis (= P. neriifolius ?), P. lophatus and P. spathoides. We present maps of these
species here, noting that they are incomplete in cases where the species would prove to be distinct
if better known, or become subsumed in other maps if they turn out to be synonyms.

Map MA-28. Podocarpus macrocarpus (black) [n=13, m=13, h=10,

1908–1979]; Podocarpus palawanensis (red) [n=1, m=1, h=2, 1984]

Podocarpus macrocarpus appears to be very similar to P.

rumphii and is possibly only a variety of it (Farjon, 2010a,
vol. 2, p. 857). The type locality, Twin Peaks, is very close to
Mt. Santo Tomas in Benguet Province, Luzon, Philippines
where David de Laubenfels, the author of this species, made
later collections. Ten localities are known, all in N Luzon,
but since De Laubenfels collected it in 1978–79 no further
specimens seem to have been identified as this species.
Those with indications of altitude record 2000–2100 m occur-
ring in ‘cloud forest’; P. rumphii occurs below 1600 m. The
data from the known collections are insufficient to establish
the altitude range of P. macrocarpus, or its distribution and
308 malesia

­habitat. Podocarpus palawanensis is only known from its type collection, C. E. Ridsdale SMHI 1502
(K, L) collected in 1984 in the Pagdanan Range, Palawan, Philippines. It is similar to P. polys-
tachyus (map MA-9 on p. 300) which has been collected from several localities on this island.
Podocarpus macrocarpus is listed as Endangered (EN) and P. palawanensis as Critically Endan-
gered (CR) on the IUCN Red List.

Map MA-29. Podocarpus
ramosii (black) [n=12, m=12,
h=10, 1886–1997]; Podocarpus
atjehensis (red) [n=6, m=5,
h=5, 1937–1975]

Podocarpus ramosii is
known from two localities
in Luzon, Philippines (Mt.
Banáhao and Mt. Isarog)
and from Gunung (mtn.)
Beratus, Kalimantan Timur,
Borneo. Ten of the twelve
collections found in herbaria are from Mt. Banáhao; the most recent collection is from Mt. Isarog.
This species was described and named P. rotundus by David de Laubenfels and renamed by Rob-
ert Mill because the name P. rotundus had been used earlier for a fossil. It may belong to P. pilgeri
which does occur in Luzon and in Borneo and is possibly a variety of it. Podocarpus atjehensis
was first described as a variety of P. neriifolius but raised to species rank by David de Laubenfels.
In Flora Malesiana 10 (3), fig. 53 De Laubenfels (1988) presented a map with two dots, one in
Aceh, N Sumatera, the other far to the east at the Danau Paniai Lakes (‘Wissel Lakes’) in Papua,
W New Guinea. Three localities are now known in N Sumatera, but the location in New Guinea
is still the only one recorded from two old collections with duplicates in K and L. The collections
in Sumatera came from “montane scrub” and ‘mossy forest’ at 2500–2900 m a.s.l. while the New
Guinea specimens were collected at 1750–1800 m at the base of a mountain in what must have
been lower montane rainforest. It is an unlikely distribution for a taxon at species rank. Without
more, fertile specimens obtained from field work it is difficult to evaluate the validity of this spe-
cies or to map its distribution.

Map MA-30. Podocarpus lophatus (black) [n=3,

m=3, h=4, 1907–1995]; Podocarpus spathoides (red)
[n=5, m=5, h=5, 1973–1978]

Podocarpus lophatus is only known from three

collections in two localities in the Philippines.
The type collection, which was the only collec-
tion De Laubenfels knew when he described it
as a new species, was taken from Mt. Tapulao in
Luzon, in 1907; the second collection from there
is undated but early 20th century. The third col-
lection is much more recent, 1995, and comes
from the summit area (2585 m) of Mt. Halcon in
Mindoro, where it is said to be very common. It
is perhaps a rare species but more material needs
to be collected and studied to establish its status
both taxonomically and biogeographically. Its conservation status is listed as Vulnerable (VU) on the
IUCN Red List. Podocarpus spathoides is only known with certainty from the type locality, Gunung
Ledang (Mt. Ophir) in Peninsular Malaysia. Another locality is possibly in Lawas, N Sarawak, Borneo,
 malesia 309

based on a collection at K which became mislaid. Previous circumscriptions of the species both mor-
phologically and geographically (De Laubenfels, 1988, followed by Farjon, 2010a) are incorrect and
collections assigned to it from the Moluccas, the Louisiades Archipelago and the Solomon Islands
belong to other species (Mill & Whiting, 2012). This too is a species still only known from a few speci-
mens which leave gaps in the description for lack of (mature) fertile material.

Dacrydium

The genus Dacrydium (Podocarpaceae) has 22 species (one of which was described as a nothospe-
cies) of which 15 occur in Malesia. It is the second most diverse genus in Podocarpaceae, but well
below Podocarpus (98 species). The genus has a tropical distribution with a southern extension
into the temperate latitudes of New Zealand; northward it does not extend beyond the tropics
although coming close in N Viet Nam. If one draws the extent of occurrence (EOO) around the
peripheral dots on the map, it becomes evident that Dacrydium is anomalously absent in Jawa
and the Lesser Sunda Islands as well as in tropical Australia, particularly coastal Queensland.
The genus appears to have become extinct in Australia during the Miocene (Florin, 1963). In
the SW Pacific it is absent in Vanuatu and the Santa Cruz Islands. Borneo has seven species and
New Guinea has eight species. Towards the outer periphery of its range the number of species
drops from four in Peninsular Malaysia to just one each in Indochina and Hainan Island (China)
and from five (1 nothospecies) in New Caledonia to a single one in New Zealand. The centre of
species diversity is in Malesia on either side of Wallace’s Line. Only two species, D. beccari and
D. xanthandrum occur both in Borneo and in New Guinea, so it appears that this biogeographi-
cal dividing line has some significance separating species in the western half of Malesia from the
eastern half. This division is used to group the maps of the species, starting with the two wide-
spread species that cross the line.

Map MA-31. The distribution of the genus Dacrydium.

310 malesia

Map MA-32. Dacrydium xanthandrum [n=57, m=54, h=16, 1884–2008]

Dacrydium xanthandrum occurs widely in Malesia, from N Sumatera, Borneo, Luzon, Mindanao,
Sulawesi, New Guinea and New Britain to the Solomon Islands. It is absent in Peninsular Malaysia
(Farjon, 2010b). In most of the large islands it has only been found locally, with more extensive
occurrences in Sabah, N Sarawak and the Huon Peninsula in Papua New Guinea. Dacrydium xan-
thandrum can grow to a large tree in lowland to montane evergreen rainforest, but also occurs in
low ‘mossy forest’ on exposed ridges. In tall forest it occurs as a sporadic emergent, often among
other large trees, while on exposed ridges it is sometimes dominant. No specifics are known that
may explain how this needle-leaved, ostensibly light demanding tree establishes and maintains
itself amongst broad-leaved, shade tolerant angiosperms, but similarities with the New Zealand
species D. cupressinum (map AU-57 on p. 380 may indicate part of the answer. On the other hand,
unlike its New Zealand congener, it can also ‘escape’ competition by establishing in situations
unfavourable to broad-leaved trees. Its altitude range is large, from near sea level in the Solomon
Islands to 2650 m on Mt. Kinabalu in Borneo.

Dacrydium xanthandrum. The photo on the left shows primary montane rainforest in the Crocker Range,
Sabah (Malaysia), Borneo. The white arrow points at the emerging crown of the conifer. The photo on the
right shows a free standing tree in the same area, due to forest clearing for a newly built road. Photo credit
Aljos Farjon.
 malesia 311

Map MA-33. Dacrydium beccarii [n=94, m=85, h=14, 1932–1996]

Dacrydium beccarii is the other widespread species in Malesia and generally more common than
D. xanthandrum. It occurs from N Sumatera and Peninsular Malaysia through mostly northern
Borneo, the Philippines, Sulawesi, the Moluccas, New Guinea and New Britain to the Solomon
Islands. Highest densities of populations are present in N Sumatera and in Malaysia (both Penin-
sular and in Borneo) where many steep mountain ridges and summits between 500 m and 2500
m are occupied by this low canopy tree, often the dominant species in ‘mossy forest’ in skeletal
soil or sphagnum peat on karst limestone or igneous rock of various types. High humidity charac-
terizes this habitat, where conifers have an ‘edge’ over most angiosperms in terms of dominance
and/or tree size and where epiphytes of all kinds from lichens and mosses to orchids and pitcher
plants (Nepenthes spp.) make up a high proportion of plant diversity. Another habitat type where
D. beccarii can be found is ‘kerangas’ forest on leached sandy soil, often covered by sphagnum
peat where runoff slows or stagnates in more or less level areas on sandstone plateaus.

Map MA-34. Dacrydium pectinatum [n=61,

m=54, h=10, 1865–1996]

Dacrydium pectinatum occurs on

Hainan Island (China), in the Philip-
pines, in Borneo (including the islands
Karimata and Serutu) and in Sumat-
era on Belitung (Billiton) Island and
on Bangka Island. In Flora Malesiana
10 (3): 364 (De Laubenfels, 1988) it is
also reported to occur on the Natuna
Islands in the South China Sea N of the
western end of Borneo, but it was not
shown on the map and we have found
no herbarium specimen to confirm this.
On occasion D. pectinatum is a tall tree
emergent above the canopy of (sub)
montane rainforest, but more often it occurs in lower, more open vegetation either in ‘kerangas’
forest or among shrubs on nutrient-poor soils or rocky ridges. The altitude range is from near sea
level to 1500 m, with trees becoming scarce above 600 m altitude. Deforestation and transforma-
tion of its habitat into oil palm plantations are thrreatening this species with extinction and it
is listed as Endangered (EN) on the IUCN Red List.
312 malesia

Map MA-35. Dacrydium elatum

[n=85, m=73, h=13, 1822–2012]

Dacrydium elatum occurs in S

China: Guangxi Province; in Indo-
china: Kampuchea [Cambodia],
Lao PDR, Thailand, Viet Nam; in
Malesia: Borneo (Brunei, Sabah
and Sarawak), Philippines, Penin-
sular Malaysia (including Penang
Island) and Sumatera (W coast).
Specimens from Hainan Island in
Chinese and other herbaria iden-
tified as D. elatum or as D. pierrei
Hickel, a synonym of D. elatum
(Roxb.) Wall. ex Loudon, are not
that species but belong to D. pec-
tinatum, the only native species
there. Only one correctly identified specimen was found on the Chinese Virtual Herbarium (CVH)
website, collected in 1966 from what is now Guantouling National Forest Park near Beihai on the
coast of Guangxi, held at IBK. Dacrydium elatum is more common and widespread in mainland
Southeast Asia than in the Philippines and in Borneo and Sumatera. In Borneo it is limited to
the northern parts; it is therefore possible that this species of Dacrydium is mainland Asian in
origin and could have spread across the Strait of Malacca and the South China Sea when these
fell largely dry during Pleistocene glacial maxima. The altitude today ranges from 250 m to 2350 m
so it could well have survived at sea level in a cooler climate.

Map MA-36. Dacrydium comosum

(black) [n=18, m=18, h=8, 1922–1990];
Dacrydium gracile (red) [n=12, m=12,
h=4, 1960–1979]

Dacrydium comosum is probably

endemic to Peninsular Malaysia.
It is known from four locations,
three in Pahang: Genting High-
lands (dominant on the ridges of
Gunung Ulu Kali), Pine tree Hill
(on ridges amid golf courses and
other development) and Gunung
Tahan, and Gunung Jerai in Kedah. It was stated to also occur in Sumatera (Farjon, 2010b) but
no herbarium specimens have been found to substantiate this. Dacrydium comosum occurs on
exposed mountain ridges as a local dominant in stunted ‘mossy forest’ on rocky acidic soil or
shallow peat; in one area at altitudes between 1170 and 1440 m a.s.l. (the altitudinal range is
incompletely known). This species is listed as Endangered (EN) on the IUCN Red List. Dacrydium
gracile occurs on Mt. Kinabalu and vicinity (Mt. Kinabalu National Park) and is known from one
locality in Sarawak, the Mujong-Merirai Protected Forest, where it was collected in 1964. It is a
relatively rare tree occurring scattered in lower montane rainforest, at altitudes between 950 m
and 1800 m. It is usually associated with the conifers Agathis borneeensis, Podocarpus laubenfelsii,
Sundacarpus amarus, Falcatifolium falciforme, Nageia wallichiana and Dacrycarpus imbricatus on
soils poor in nutrients (‘kerangas’ forest); in Sarawak it occurs in low canopy ‘heath’ forest on
sandstone. Dacrydium gracile is listed as Near Threatened (NT) on the IUCN Red List.
 malesia 313

Dacrydium comosum on a
foggy ridge of Gunung Ulu
Kali, Peninsular Malaysia.
Nearly perpetual clouds swirl-
ing around these mountain
ridges bring high humidity,
creating a low ‘mossy forest’
characterized by these coni-
fers and other shrubs and
numerous epiphytes from
lichens and mosses to orchids
and pitcher plants. Photo
credit Aljos Farjon.

Map MA-37. Dacrydium medium

(black) [n=28, m=28, h=5, 1934–1978];
Dacrydium ericoides (red) [n=11, m=11,
h=5, 1923–1990]

Dacrydium medium occurs in the

central part of Peninsular Malay-
sia and in Aceh, N Sumatera. This
species grows almost exclusively
on steep ridges and summits at
altitudes between 1050 and 1800
m in Peninsular Malaysia and at
2500–2600 m in Sumatera. Mostly
shrubs, occasionally small trees
when growing a little ways down
slope among taller trees, this spe-
cies is relatively common but has only a small range. The two Sumatran localities may not be
the only ones, but we have found no more than two herbarium collections (both in L), the most
recent of which was collected in 1975. Dacrydium ericoides has a limited distribution in Sarawak,
Borneo, where it is known from six localities. This too is a species of mountain ridges and sum-
mits, collected from 1020 m to 2200 m a.s.l. where it is locally common in ‘mossy forest’. It can
occasionally grow to a tree 18 m tall and with 30 cm d.b.h. when located off the ridge in a shel-
tered spot. Dacrydium medium is listed as Vulnerable (VU) on the IUCN Red List.

Map MA-38. Dacrydium spathoides

[n=3, m=3, h=5, 1939]

This poorly known species is rep-

resented by three herbarium col-
lections, made in the same locality
along an altitudinal ‘transect’
following the upper Idenburg
River in the Maoke Range, Papua
(W New Guinea). These specimens
are from “18 km SW of Bernhard
Camp” (‘Top Camp’); all were collected by L. J. Brass in February 1939 and several duplicates are kept
at A, BRI, K, L and LAE. That first gathering was undertaken during a Dutch-American ­expedition
314 malesia

into then uncharted territory in the interior mountains of New Guinea and the effort has not been
repeated. Dacrydium spathoides was observed to be a canopy tree in evergreen ‘mossy forest’ at
2150–2200 m altitude. Its description as a new species by David de Laubenfels came 30 years after
its discovery, based on these few herbarium specimens only. This is not much to go by taxonomi-
cally or biogeographically and this taxon must wait until the area is revisited and new samples are
gathered before we can decide on its status and distribution.

Map MA-39. Dacrydium cornwal-

lianum [n=16, m=16, h=5, 1912–1991]

Dacrydium cornwallianum is
endemic to the island of New
Guinea. It occurs in only a few
areas, most commonly in the area
of the Danau Paniai Lakes (‘Wissel
Lakes’) in Papua (W New Guinea)
and in the Mendi Subdistrict of the
Southern Highlands, Papua New
Guinea. Its most common habitat
is edges of swamps and bogs, where it can form almost pure stands. This species is also found on
mountain ridges in ‘mossy forest’ but there it is uncommon, and sporadically as a canopy tree mixed
with angiosperms, probably Nothofagus (not mentioned with herbarium collections). In these cir-
cumstances it may attain 30 m, but on the ‘black peat’ it is of much more modest stature.

Map MA-40. Dacrydium novoguineense [n=32, m=31, h=8, 1912–1994]

Dacrydium novoguineense is, despite its species name, not endemic to New Guinea, but also
occurs in the Moluccas and in Sulawesi. The type was collected in the Arfak Mountains of the
Birds Head Peninsula, Papua (W New Guinea) in 1913 (L. S. Gibbs 5648, BM, K) which is in dis-
tance terms about midwway of its range. However, if the number of herbarium collections and
their localities reflect relative abundance, this species is more common in New Guinea than in
the islands to the west, with only five mapped localities. It can indeed be abundant to dominant
in the canopy on mountain ridges. On lower slopes it is an emergent tree and can attain 30 m
and some observations noted on herbarium labels seem to indicate that it regenerates quickly
after forest fires. The altitude range of this species is very large, from 50 m to 3000 m according
to data from herbarium labels.
 malesia 315

Map MA-41. Dacrydium nidulum [n=60, m=59, h=8, 1855–1998]

Dacrydium nidulum has a similar distribution in Malesia as D. novoguineense, from Sulawesi in the
west to the Louisiades Archipelago in the east, but in addition occurs far to the east of this area
in Fiji. This disjunction in its range seems at first sight an anomaly, but its presence on islands
beyond the far eastern end of New Guinea could indicate an answer to how this came about.
Although no specific ‘mutualisms’ are known in the literature, seed dispersal must be similar to
that of Podocarpus species, but seeds and coloured, succulent receptacles of Dacrydium species
are much smaller. This could imply small passerine birds on migratory routes through the island
chains of Malesia and the SW Pacific. Dispersal to a remote island would be a chance event, but
once a species is established, further dispersal on that island and to neighbouring islands becomes
more likely to occur; indeed it may become inevitable over time. In this manner D. nidulum may
have arrived in Fiji, bypassing all the islands that are situated between the Louisiades and Fiji.
Whether it also had arrived on the island of Seram, one of the Lesser Sunda Islands, as indicated
in Flora Malesiana 10 (3): 365 (De Laubenfels, 1988) seems doubtful. No herbarium specimen has
been found in L or elsewhere to substantiate the dot on the map in fig. 13 of that account; the
only conifer species vouchered by herbarium specimens occurring on this island are Dacrycarpus
imbricatus and Sundacarpus amarus. For this reason we interpret this as an error, possibly a mis-
identification. Dacrydium nidulum has been found as a canopy tree in lowland tropical rainforest,
as well as in peat bogs at 2300 m a.s.l.

Map MA-42. Dacrydium leptophyllum (black) [n=1, m=1, h=2, 1911]; Dacrydium magnum (red) [n=8, m=8,
h=6, 1937–2008]
316 malesia

Dacrydium leptophyllum is only known with certainty from the type, collected in 1911 by A. C.
de Kock on Mt. Goliath in Papua (W New Guinea) and kept in BO, with an isotype (duplicate
specimen, in this case a mere fragment) in S. As a result, this species is poorly known and more
collecting needs to be done to evaluate it. Perhaps it is not distinct, as mature fertile specimens
were not collected. Dacrydium magnum is known from a few more specimens, but its known dis-
tribution from these is decidedly odd. This species, described and named by David de Laubenfels
in 1969 from Obi Island in the Moluccas, has also been collected on Tagula Island (Louisiades
Archipelago), on Normanby Island (D’Entrecasteaux Islands) and on Choiseul (Solomon Islands).
In Flora Malesiana 10 (3): 368 (De Laubenfels, 1988) Guadalcanal and Santa Ysabel in the Solomon
Islands are also mentioned, but no herbarium specimens could be found. No specimens from New
Guinea are known and a few labelled as this species turned out to be misidentified. If dispersal by
birds is largely responsible for the island distribution of this species, it is difficult to see how New
Guinea could be missed. Either the distribution is very incompletely known, or the specimens on
either side of New Guinea belong to different species. The collection L. J. Brass 27821 (A, K, L, S, US)
is the type of Dacrydium beccarii var. rudens, now treated as a synonym of D. magnum (De
Laubenfels, 1988; Farjon, 2010a) and was collected on Mt. Riu on Tagula Island. We may have to
revisit this taxonomy. Meanwhile this species is listed as Vulnerable (VU) on the IUCN Red List.

Dacrycarpus

The genus Dacrycarpus (Podocarpaceae) has nine species, of which seven occur in Malesia. One
species is endemic in New Zealand (D. dacrydioides, map AU-58 on p. 381) and D. vieillardii (map
AU-99 on p. 409) is found in New Caledonia. The genus is distributed from N Myanmar [Burma]
and S China to Fiji and New Zealand, similar to Dacrydium, but also present in Jawa and the
Lesser Sunda Islands. A gap is formed by its absence in the Solomon Islands and, like Dacrydium,
it is not present in Australia. Five species occur in New Guinea and five in Borneo. Only one spe-
cies, D. inbricatus, is very widespread. Although some species do not cross Wallace’s Line, several
others do and there is not such a clear division between east and west as in Dacrydium. A com-
prehensive phylogenetic analysis which could assist in explaing the distribution of the genus was
not available at the time of writing.

Map MA-43. The distribution of the genus Dacrycarpus.

 malesia 317

Map MA-44. Dacrycarpus
imbricatus [n=223, m=204, h=19,
1810–2009]

In this very widespread spe-

cies, 3–4 varieties are recog-
nized, but the distinctions
are small and one, var. patu-
lus, was found to be incon-
sistent and has been sunk
altogether (Farjon, 2010a,
2010b). Here we map them
together as the species. With
the exception of New Cale-
donia and New Zealand, the
distribution of Dacrycarpus imbricatus virtually equals that of the genus. Only in New Guinea are
other species in the genus taken together more common than this species. In mainland SE Asia, in
the Greater and Lesser Sunda Islands, in Vanuatu and in Fiji D. imbricatus is the only species pres-
ent. The distribution of this species is most similar to that of Podocarpus neriifolius (map MA-7
on p. 298) and differs in that it does not extend as far N into China, is very rare in the Moluccas
and absent in the Solomon Islands, but is present in the Lesser Sunda Islands. What this could
indicate is that both species are highly successful dispersers, but that D. imbricatus is less tolerant
of low winter temperatures which can occur in the mountains of interior China. Absences in most
of the Moluccas and in the Solomon Islands, as well as islands in the Indian Ocean off Sumatera
and the Andaman Islands, are probably accidents of (lack of) dispersal, as this is always a chance
event. Dacrycarpus imbricatus is a tall canopy tree or emergent in tropical evergreen rainforest,
usually montane but occasionally found in the lowlands; its altitude range is very large, 200–3720
m, but more commonly upwards from 1200 m, the limit of dipterocarp rainforest, to 2500 m
beyond which a more stunted, ‘mossy’ forest type prevails. These forests are usually dominated
by Fagaceae (in New Guinea also Nothofagaceae) and often have a scattered presence of conifers,
of which Agathis, Podocarpus, Dacrydium, Dacrycarpus, Nageia, Sundacarpus and Phyllocladus are
the most common genera.

Dacrycarpus imbricatus in rainforest on the Mesilau River,

Mt. Kinabalu National Park, Sabah (Malaysia) Borneo. This
conifer occurs as scattered individual trees which eventu-
ally emerge above a canopy of large-leaved angiosperms.
There are many seedlings and some saplings on the forest
floor, but it seems that only very few succeed to maturity.
Photo credit Aljos Farjon.
318 malesia

Map MA-45. Dacrycarpus cumingii [n=40,

m=40, h=20, 1841–1992]

Dacrycarpus cumingii is mostly distrib-

uted in the Philippines, where it occurs
in Luzon, Mindanao, Negros and on
Palawan Island. A disjunct locality is
Mt. Penrissen in westernmost Sarawak;
there are also two collections by Van
Steenis at BO from Aceh, N Sumatera.
These localities are probably true outli-
ers for this high altitude species, which
occurs usually from 2000 m to 2700 m
but up to 3314 m on Gunung Kemiri in
Aceh, Sumatera. The intervening high country in Borneo has been well collected and it has
not been found there apart from this one mountain. The habitat is described as ‘mossy forest’
or primary forest to 25 m tall, so it can occur in two types, but both are characterised by poor
soils, the first on steep ridges and summits, the second on leached sands in more level terrain.
In both situations conifers and small-leaved angiosperms (Casuarinaceae, Myrtaceae) exist in
mixture, but large-leaved angiosperms are mostly absent in the canopy.

Map MA-46. Dacrycarpus compactus [n=54,

m=54, h=12, 1911–1998]

Dacrycarpus compactus is endemic to

New Guinea, where it occurs along the
central mountain chain. A peculiar gap
in its distribution exists in the Western
Highlands of Papua New Guinea, or
between the Sirius Ridge in the east and
the Star Mountains near the Indonesian
border in the west. This gap is filled by other species in the genus, mainly by Dacrycarpus expan-
sus. Its distribution then continues westwards as far as Mt. Jaya, itself an isolated locality for this
species. Dacrycarpus compactus is a highland species occurring in subalpine shrubberies and on
the fringes of alpine tussock grassland dominated by Deschampsia klossii. It is common in conif-
erous high montane forest with Papuacedrus papuana, Podocarpus spp., and a few angiosperms
and then becomes more abundant and often a dominant emergent tree in mossy low forest and
shrubbery on rocky terrain fringing wet tussock grasslands on peat. The altitudinal range of this
species is 2800–4300 m a.s.l.

Map MA-47. Dacrycarpus expansus [n=17,

m=17, h=7, 1938–1975]

This is another species endemic to New

Guinea. Its main distribution is in the
central highlands of Papua New Guinea,
but two outlying localities are known
from herbarium collections. The one to
the E of the main area is in the moun-
tains above Goodenough Bay; the one to
the W is near Lake Habbema in W New Guinea (Papua, Indonesia). It is likely that this species
 malesia 319

occurs in a few localities between these outlying populations, but the species is relatively rare
and only a small number of herbarium collections have been made, some from the same locality
as others. Dacrycarpus expansus occurs in lower montane to high montane forests (ca. 2000 m to
3500 m a.s.l.), often on the margins of tussock grassland with tree ferns (Cyathea). It can be mixed
with Papuacedrus papuana or occur in nearly pure stands. The soil is often water-logged, peaty
and acidic.

Map MA-48. Dacrycarpus cinctus [n=44, m=44, h=10, 1902–2000]

Dacrycarpus cinctus occurs in New Guinea, the Moluccas, Sulawesi and Borneo. It is most com-
mon in the highlands of Papua New Guinea and in the mountains of central Sulawesi; more scat-
tered occurrences are known from the Maoke Range in Papua (West New Guinea), the island of
Seram in the Moluccas and one locality in Borneo (Gunung Mulu in Sarawak). In New Guinea
Dacrycarpus cinctus is another highland species occurring in high montane to subalpine habitat,
at altitudes between 1800 m and 3600 m, in evergreen tropical rainforest at the lower range to
‘mossy’ forest and margins of tree-fern grassland at the higher altitudes. In Sulawesi this species
has been found at 900 m a.s.l., but other collections range from 1700–3000 m and therefore indi-
cate a similar habitat. More collecting is probably needed to complete the map of this species,
which probably occurs on other high mountains within its range.

Map MA-49. Dacrycarpus steupii [n=36, m=34, h=6, 1937–1980]

320 malesia

Dacrycarpus steupii is most common and widespread in New Guinea, where it follows the cen-
tral mountain chain. More isolated occurrences are in the Bird’s Head Peninsula of New Guinea,
Sulawesi and Borneo. In Sulawesi Dacrycarpus steupii is still common in the Latimodjong Mountains;
another location is Mt. Roroka Timbu in N-Central Sulawesi. In Borneo, it may now be extinct on the
E coast, where it was collected on Gunung Beratus near Balikpapan in 1952 but was not found
again in the early 1980s due to deforestation. A more recent specimen (1980) was collected in Kali-
mantan Barat near Kembajan (G. Shea 275022 at L). Dacrycarpus steupii is common in mossy forest
and subalpine shrubberies from montane to alpine zones. Its altitude range is between 860 m and
3470 m and at lower elevations it can grow to a large tree, subject to logging. It is these lower locali-
ties that are most threatened with deforestation. Because it is still widespread and common in New
Guinea the global conservation status of Dacrycarpus steupii is outside a threatened categorry of
the IUCN Red List, but in Borneo and N-Central Sulawesi the situation is more serious.

Falcatifolium

The genus Falcatifolium (Podocarpaceae) has six species, of which five occur in Malesia and one
in New Caledonia (F. taxoides, see map AU-96 on p. 407). In Malesia, the genus is widespread
from Peninsular Malaysia to Papua New Guinea and occurs in the Malay Peninsula, the Riau
Islands, Borneo, Philippines, Sulawesi, Obi Island in the Moluccas, and New Guinea. The genus is
absent in the Greater and Lesser Sunda Islands, the Bismarck Archipelago and all islands between
New Guinea and New Caledonia, so that F. taxoides is disjunct from the remainder of the genus.
Is its presence in New Caledonia an ‘accident’ of long distance dispersal from Malesia? We need
to know what its phylogenetic relationship is with the other five species. Biffin et al. (in Turner &
Cernusak, eds. 2011) included three species in their molecular analysis and F. taxoides was sister
to F. falciforme and F. gruezoi. If this topology is upheld when all six species are included, it would
indicate that Falcatifolium has a Gondwanan origin and spread into Malesia where it diversified
into several species. Falcatifolium taxoides could then be interpreted as a relict from an ances-
tral lineage, not the result of long distance dispersal from Malesia. The fossil record gives some
support for this hypothesis because the genus has been identified from the Eocene of Victoria,
Australia (Hill & Brodribb, 1999).

Map MA-50. Distribution of the genus Falcatifolium.

 malesia 321

Map MA-51. Falcatifolium
angustum [n=4, m=4, h=4,
1959–1966]

Falcatifolium angustum is
a poorly known species,
collected between 1959
and 1966 in two localities
on the coast of Sarawak,
Borneo. The northernmost
locality, where the type
specimen was collected, is
in Niah National Park, the
southern locality is in Bako
National Park. These herbarium collections were identified and described as a new species in 1969
and nothing that was or could be identified as this species has been collected since. The descrip-
tion was based on male specimens and the seed cones were and remain unknown. Falcatifolium
angustum occurs in ‘kerangas’ (forest on podzolised white sands) at 90–240 m altitude near the
coast. The two populations occur in open forest with among other trees Gymnostoma sp. (Casu-
arinaceae), Parastemon sp. (Chrysobalanaceae) and Shorea albida (Dipterocarpaceae). New col-
lections need to be made to establish its validity as a distinct species and if it is distinct, where it
still occurs. The coast of Sarawak is rapidly being deforested and if it occurred in other localities,
it may now have disappeared there. We do not even know if it still exists where it was found.
Based on its two known localities in ‘protected’ areas this species has been listed as Endangered
(EN) on the IUCN Red List.

Map MA-52. Falcatifolium
papuanum (black) [n=21,
m=20, h=6, 1923–1981]; F.
sleumeri (red) [n=1, m=1,
h=1, 1961–1961]

These two species are

endemic to New Guinea.
Falcatifolium papuanum
occurs in the highlands of
Papua New Guinea, from
the border with Indonesia
(Papua) to the Owen Stan-
ley Range. Most herbarium collections were made in the Morobe Province (Wau Subdistrict and
Mt. Kaindi). It is usually a canopy or subcanopy tree locally common in montane rainforest domi-
nated by Fagaceae (Lithocarpus), Nothofagaceae, Myrtaceae, Cunoniaceae, and Podocarpaceae.
The altitude range is from 1300 m to 2300 m a.s.l. Falcatifolium sleumeri is only known from the
type specimen, collected in 1961 by Pieter van Royen on Mt. Nettoti in the Arfak Mountains of the
Bird’s Head Peninsula of W New Guinea (Papua, Indonesia). It is apparently a low, decumbent
shrub growing in “podocarp-Myrtaceae forest” or “dark mossy forest” at 1920 m a.s.l. which is on
the summit ridge or crest. No herbarium specimens have been collected since and the specimen
from which this species was named and described was sterile. The locality is remote (no road
access) but needs to be revisited by botanists.
322 malesia

Map MA-53. Falcatifolium falciforme (black) [n=63, m=61, h=6, 1865–1994]; Falcatifolium falciforme trees on
F. gruezoi (red) [n=15, m=14, h=8, 1906–2009] a mountain ridge in the Crocker
Range, Sabah, Borneo. Photo
credit Aljos Farjon.

Falcatifolium falciforme is the most widespread and common species of the genus in Malesia. It
occurs in Peninsular Malaysia, on Lingga Island in the Riau Group, and in Borneo. This species
is most commonly found in low ‘mossy’ forest on mountain ridges, but occurs in a wide range of
habitats, from coastal ‘kerangas’ to montane rainforest. The altitude range is consequently wide,
from 300 m to 2100 m. In the rainforest it is uncommon and usually limited to a few scattered
individual trees. Seedlings can be abundant under tall trees, but seldom grow to mature trees
which may occasionally attain 40 m. For this to happen, it is likely that canopy removal as a result
of a major forest disturbance is necessary. On mountain ridges competition from angiosperms
is less severe. Falcatifolium gruezoi occurs in the Philippines on the islands of Luzon, Mindoro,
Panay and Mindanao. The locations Davao and Surigao (minor areas in Mindanao) cited by Mer-
rill (1934) under Dacrydium falciforme and by De Laubenfels (1988) from that earlier source under
F. gruezoi have not been traced to herbarium collections and are not shown on the map. Possibly
three of the five collections cited by Merrill come from Mindanao but it is not clear which, if
any. In Sulawesi the localities found with herbarium specimens for this Atlas only agree with one
of three mentioned in Flora Malesiana 10 (3): 373 (De Laubenfels, 1988). Both datasets agree on
the occurrence on Obi Island in the Moluccas. It seems clear that the distribution of this species
remains imperfectly known. Falcatifolium gruezoi occurs in lower montane to montane rainfor-
est. It is commonly associated with other conifers, e.g. Agathis dammara, Dacrydium spp., Nageia
wallichiana, and Sundacarpus amarus. Angiosperms are often abundant but not dominant.

Nageia

The genus Nageia (Podocarpaceae) has five species, three of which occur in Malesia and are
presented in this chapter. The two species outside the region are N. nagi and N. fleuryi, both are
treated in the chapter Mainland Asia and Japan. The distribution of the genus extends from the
Western Ghats in S India to the D’Entrecasteaux Islands in the Solomon Sea and from Honshu
in Japan to the Lesser Sunda Islands. It is present throughout S China including Taiwan and
Hainan, in Indochina and W to Meghalaya in NE India, down the length of the Malay Peninsula
and in most of Malesia. In Jawa it becomes rare, with a presence only in W Jawa (Jawa Barat) and
the genus is absent from most of the Lesser Sunda Islands, with only two localities known from
herbarium collections. It is also absent in the Bismarck Archipelago and the Solomon Islands, so
it reaches its easternmost location on Normanby Island off the eastern tip of New Guinea. The
 malesia 323

occurrence in the Western Ghats is not only a disjunct locality, it is the only natural occurrence
of a conifer in the whole of the Indian subcontinent (not in India, which extends into the Hima-
layas beyond the subcontinent proper). It is likely the result of a westward dispersal event that
occurred from a population of N. wallichiana within its wide range to the east. The northernmost
outliers in Honshu are around 35° 20’ N and at lowland altitudes (below 200 m); climate is the
limiting factor to northern expansion. Not much is known yet about relationships among the
species; only the more common three species were sampled in the study on podocarp phylogeny
by Biffin et al. (in Turner & Cernusak, eds. 2011).

Map MA-54. The distribution of the genus Nageia.

Map MA-55. Nageia wallichiana [n=193, m=154, h=22, 1830–2009]

324 malesia

Nageia wallichiana is the most widely distributed species in the genus. With the exclusion of nearly
all of China and of Japan, its distribution virtually coincides with that of the genus and includes
the western outliers in India as well as the easternmost occurrence on Normanby Island. This is
one of the most extensive ranges known in conifers and comparable to Dacrycarpus imbricatus
(map MA-44 on p. 317) and Podocarpus neriifolius (map MA-7 on p. 298) with much the same
distribution. Dispersal success undoubtedly is the basis for this pattern, but why exactly these
species have been so successful in spreading while their congeners in the same region have much
more limited distributions remains largely unanswered (see e.g. Enright & Jaffré in Turner & Cer-
nusak, eds. 2011). More detailed ecological studies of these three podocarps are required, such as
investigations into which animals are primarily responsible for seed dispersal and how effective
they are for these species in comparison to related species. Nageia wallichiana is a strictly tropi-
cal conifer, occurring from lowland forest near sea level, including dipterocarp forest, to montane
forest at around 2100 m. It can reach the canopy to 50 m, but is not a long-lived tree and prob-
ably emerges in small forest gaps (not wider than canopy height) from randomly dispersed seeds.
Unlike Agathis it is not gregarious, indicating a lesser dependence on large scale disturbances.
Both are truly broad-leaved conifers, in fact their leaves are often hard to distinguish. In ‘kerangas’
forest N. wallichiana can be more abundant than in tall closed-canopy rainforest.

Map MA-56. Nageia motleyi [n=39,

m=30, h=9, 1857–1961]

Nageia motleyi occurs in the western

part of Malesia, in the southern part
of the Malay Peninsula, including
southernmost Thailand (no herbar-
ium records found) and Peninsular
Malaysia, in Sumatera and Borneo.
In all of these areas this species is
rare and only known from widely
separated localities (perhaps with
some concentration in the western parts of Peninsular Malaysia). Nageia motley is a lowland
species, known from ca. 15 m to 500 m a.s.l. as a rare constituent of “bindang-dipterocarp” forest
in Sarawak, but it is more often found in less tall evergreen forests on podzolic sands, where it is
scattered among numerous angiosperm tree species and occasional conifers. It apparently regen-
erates after disturbance and is also found in secondary forest. Both the geographical range and
the habitat of N. motley partly overlap with N. wallichiana and there may be issues with correct
identification. The herbarium collections used for this map have all been seen and verified by
specialists. Nageia motleyi is listed as Vulnerable (VU) on the IUCN Red List.

Map MA-57. Nageia maxima [n=7, m=7, h=5, 1961–1973]

This species remains in some respects poorly known.

The “few collections” De Laubenfels (1988) knew
(three were his own) have not been added to since
1973 and the two known localities, Bako National
Park and Naman Forest Reserve in Sarawak, Malay-
sia have not been added to either. Nageia maxima
has exceptionally large leaves (up to 34 cm long and
9.5 cm wide) but remains a shrub or stunted tree to
10 m tall, as far as known. It occurs in peat swamps
but also on low ridges on dry ground, from near sea
level to 120 m. Despite its occurrence in protected
 malesia 325

areas it is listed as Endangered (EN) on the IUCN Red List, in part because its habitat is being
destroyed on a large scale by expanding oil palm plantations between the two known localities.

Retrophyllum

Map MA-58. Retrophyllum vitiense (Podocarpaceae) [n=41, m=40, h=9, 1860–1994]

Retrophyllum vitiense has an extensive but scattered distribution from Fiji, where it was first dis-
covered, Vanuatu (Banks Islands), Santa Cruz Islands, New Ireland, New Britain, New Guinea
to the Moluccas (Morotai). It has not been found in the main Solomon Islands, which seems to
be the most obvious gap. Limited or uneven collecting efforts on some islands, e.g. in Vanuatu
(Frodin, 2001: 892) may be partly responsible, but this seems less likely for the Solomon Islands,
where it has been found in the Santa Cruz Islands, a distant group of small islands belonging to
this state. Retrophyllum vitiense is a tall tree of tropical lowland to montane rainforest, usually
occurring as scattered individual emergents. Its altitude range is from near sea level to 1800 m;
on the islands in the Pacific Ocean it is moslty a lowland tree, in New Guinea and the Bismarck
Archipelago it usually occurs above 1100 m.

Sundacarpus

The genus Sundacarpus (Podocarpaceae) has a single species, Sundacapus amarus; it is closely
related to Prumnopitys which has a disjunct distribution in South America and the SW Pacific
(map SA-5 on p. 432). The two genera meet in a small area in N Queensland. In some phylogenetic
analyses based on molecular data (e.g. Biffin et al. in Turner & Cernusak, eds. 2011) Sundacarpus
amarus appears ‘nested’ within Prumnopitys, which could be an argument to merge it with that
genus. The approach followed here (Farjon, 2010a) is to recognize morphological distinction and
accept the hypothesis that Sundacarpus derived from within Prumnopitys. Given the common
area of occurrence in N Queensland, it could be assumed that Sundacarpus had its origin there.
However, from the (fragmentary) fossil record we know that Prumnopitys has existed in Tasmania
(fossils of Sundacarpus are not known). There is the likelihood that the current distribution in
Australia of both is a mere relict caused by the aridification of the continent. If this is correct,
Sundacarpus had, like other southern podocarps, a Gondwanan origin and spread out into Male-
sia more recently.
326 malesia

Map MA-59. Sundacarpus amarus [n=111, m=107, h=16, 1888–2003]

The distribution of Sundacarpus amarus is unusual for a widespread Malesian podocarp tree. It
is found in two small areas in N Queensland, the most extensive being around Cairns. Next it is
present in New Guinea as well as in New Britain (W-end of the island), New Ireland (Hans Meyer
Range) and Woodlark Island. In New Guinea it is abundant in the highlands of Papua New Guinea,
but in W New Guinea (Papua) it is only frequent in the mountains of the Bird’s Head Peninsula.
A more scattered distribution is also found in the Moluccas, Sulawesi and most of Borneo and
the Philippines; in the latter the only ‘hotspots’ are Mt. Kinabalu and vicinity and the Cordillera
Central in Luzon. Abundance recurs in a long line from the western Lesser Sunda Islands (from
Flores westward) to N Sumatera. Apart from Mt. Kinabalu, Sundacarpus amarus is extremely
rare in Borneo. Unlike other widespread species in Podocarpaceae, this species is completely
absent on the Malay Peninsula and in Indochina. This pattern of distribution is in some regions a
reversal of that seen in other Malesian conifers, which are common from the Malay Peninsula or
Borneo to New Guinea but rare or absent in the Sunda Islands (e.g. Phyllocladus hypophyllus map
MA-64 on p. 330, Podocarpus polystachyus map MA-9 on p. 300). How is this anomalous distribu-
tion to be explained? Sundacarpus amarus is rare at sea level, but becomes common at 500–2200
m so it is mainly a lower montane tree, sometimes of large size. It grows often in latosols derived
from andesite, basalt, or granite, rarely in sandy soils or in marshes. This could explain the near
absence in much of Borneo, an ancient continental fragment which experienced no volcanism
during the last 10–12 My (Hall, 2009a). Similar geologically young igneous formations predomi-
nantly form the mountains of New Guinea. The abundant occurrence in the Sunda Islands could
be related to the string of volcanoes, which continues eastward beyond Flores where rainfall
becomes too low for this wet tropical rainforest species. This volcanic arc is associated with the
subduction zone in the Java and Sunda Trenches and began as a string of small islands; only since
the Pliocene were they increasingly situated on rising larger islands (Hall, 2009b). The basalt and
andesite resulting from volcanism produce the most prevalent soil types on which this species
grows. Volcanism is also widespread in the Philippines, but Sundacarpus amarus is only common
in N Luzon, so there is certainly no direct relation with volcanoes. Chance dispersal by birds may
have taken this species to some remote localities, such as Woodlark Island and the delta of the
Mahakam River in E Borneo.
 malesia 327

Sundacarpus amarus in rainforest at Lake Barrine, Queensland, Australia. Photo credit Aljos Farjon.

Endemic Species of Mt. Kinabalu

Mt. Kinabalu [Sabah (Malaysia), Borneo] is the highest mountain between the eastern end of
the Himalayas/NW Yunnan and New Guinea. It is a solitary mountain with a summit at 4095 m.
It originated as a granitic batholith emplaced into older sedimentary rock, the centre of which
was then rapidly uplifted to form the mountain. The sediments were stripped off from the higher
parts by erosion, exposing the granite, which was carved up by glaciers during successive glacial
maxima in the Pleistocene. Lower on the mountain extensive areas are formed by ultramafic rock,
mostly serpentine; there is also sandstone belonging to the Crocker Range Formation. Beaman
et al. (1998–2004) estimated a total of ca. 5000 species of vascular plants for a 700 km² area. The
level of endemism is unknown but probably high. The mountain and its immediate surroundings,
including spur ridges and upper sections of river valleys, make up Mount Kinabalu National Park.
Although visited by many tourists the mountain is relatively well protected and offers a unique
opportunity to experience the transition from lower montane to alpine vegetation in the Male-
sian tropics along a well maintained path that leads to the summit from the end of an access road.
Conifers constitute an important part of the flora with 19 species occurring within the national
park and are present at all altitudes along this route. Four species (all in Podocarpaceae) are
endemic to Mt. Kinabalu and these are treated here together.

Mt. Kinabalu

Mount Kinabalu seen from Park Headquarters. Photo credit Richard Swann.
328 malesia

Map MA-60. Dacrydium gibbsiae [n=21, m=20, h=4,

1910–1984]

Dacrydium gibbsiae occurs on Mt. Kinabalu from

swampy ‘mossy forest’ at 1500–2000 m a.s.l. to
high rocky ridges and slopes up to 3600 m. This
species is mostly restricted to serpentine rock
and its ultramafic erosion products, but high on
the mountain it grows on granite. At lower alti-
tudes it becomes a small tree to 12 m but on the
granite it remains dwarfed to 2 m tall. Its bright
green foliage is conspicuous among the more
subdued hues of the general vegetation.

Map MA-61. Podocarpus brevifolius [n=19, m=19, h=6,

1858–1995]

Podocarpus brevifolius is locally common in upper

montane to subalpine dwarf forest on Mt. Kina-
balu and other mountain ridges and heights in the
vicinity. It grows most commonly on ultramafic
substrate and also on granite high on the moun-
tain. It has been found on ultramafic rock at 1350–
1450 m near a copper mine and on the Bambangan
Ridge at 1900 m in lower montane forest. Its alti-
tudinal range on Mt. Kinabalu proper is between
2100 m and 3800 m, where it is a constituent of a
dwarfed forest to 10 m tall, dominated by the coni-
fers Phyllocladus hypophyllus and Dacrycarpus kin-
abaluensis and the umbrella-crowned angiosperm
tree Leptospermum recurvum (Myrtaceae). Podocarpus brevifolius is the highest occurring conifer
on Mt. Kinabalu, close to the limits of vegetation, where plants are clinging to gullies and crevices
among vast expanses of bare granite forming the summit domes and peaks of the mountain.

Map MA-62. Podocarpus gibbsiae [n=8, m=8, h=5,

1910–1979]

Podocarpus gibbsiae occurs on Mt. Kinabalu and

a nearby high point, Bukit Ampuan. The largest
population is on Mt. Kinabalu. It occurs mostly
on mountain ridges between 1200 m and 2400 m
in ‘mossy forest’ on ultramafic rocks. The canopy
is lower than 25 m and often more or less open,
interspersed with shrubs in rocky areas. Podocar-
pus gibbsiae is a small to medium-size tree form-
ing part of the canopy, though never dominant
or abundant. Other common conifers in this for-
est type on Mt. Kinabalu are Phyllocladus hypo-
phyllus and Dacrydium gibbsiae, sometimes also
 malesia 329

Dacrycarpus imbricatus, the latter more often a constituent of closed canopy rainforest. Only the
trees within Mt. Kinabalu National Park are protected and this species is listed as Vulnerable (VU)
on the IUCN Red List.

Map MA-63. Dacrycarpus kinabaluensis [n=17, m=17,

h=2, 1892–1984]

Dacrycarpus kinabaluensis is a shrubby tree

restricted to the upper montane forest and sub-
alpine dwarf forest on Mt. Kinabalu. It occurs
on this mountain from ca. 2600 m up to the
tree line at ca. 3500 m a.s.l. It is growing pre-
dominantly on ultramafic rock but becomes
one of the dominant shrubs above 3000 m on
granite. It can form dense, nearly pure stands
but is commonly associated with other coni-
fers, e.g. Dacrydium gracile, D. gibbsiae, Phyllo-
cladus hypophyllus and Podocarpus brevifolius.
Common angiosperms are Leptospermum and
Rhododendron. The nutrient deficiency of rock
and soil is indicated by the common presence
of pitcher plants (Nepenthes).

Dacrycarpus kinabaluensis (shrubs in centre of picture) at 3400 m on Mt. Kinabalu. Photo credit Aljos
Farjon.

Phyllocladus

The genus Phyllocladus (Phyllocladaceae) has four species and occurs in Tasmania ­(Australia),
New Zealand and Malesia (map AU-64 on p. 384). There is only a single species in Malesia.
330 malesia

Map MA-64. Phyllocladus hypophyllus [n=137, m=134, h=13, 1851–2008]

This is the most widespread species of the genus, occurring in Borneo, the Philippines, Sulawesi,
the Moluccas and New Guinea. Phyllocladus hypophyllus is absent in Peninsular Malesia and the
Greater and Lesser Sunda Islands. There is evidence in the fossil pollen record (Morley in Turner
& Cernusak, 2011) that it slowly spread from New Guinea west- and northward across the islands
and across Wallace’s Line until it reached the eastern edge of the Sunda Shelf off W Borneo.
It may have been prevented from ‘island hopping’ along the Lesser Sunda Islands to Jawa and
Sumatera by the drier climate on these eastern islands, while it missed opportunities to cross the
Java Sea (part of the Sunda Shelf) during periods in the Pleistocene when this sea fell largely dry.
Phyllocladus hypophyllus is at present mostly a montane to high montane tree, but it does occur
in ‘kerangas’ forest on white sand down to ca. 300 m. This species could well have extended to
sea level during cooler periods. The altitude range of this widepread species is extreme, with
occurrences on Mt. Kinabalu to 3400–3500 m and in New Guinea to nearly 4000 m. Its habit con-
sequently ranges from a tall forest tree to a compact shrub. High altitude populations often have
glaucous foliage, as observed on Mt. Kinabalu.

Araucariaceae

The Araucariaceae are represented in Malesia by two genera, Araucaria and Agathis. The
genus Araucaria is in Malesia limited to New Guinea, but Agathis is widespread from Penin-
sular Malaysia to Papua New Guinea. Of the two species of Araucaria indigenous to New
Guinea, A. cunninghamii (+ var. papuana) is mapped and discussed in the chapter Australasia
(map AU-10 on p. 350). The genus Agathis is most widespread and diverse in Malesia and is
mapped and discussed in this chapter.
 malesia 331

Araucaria

Map MA-65. Araucaria hun-

steinii [n=25, m=24, h=14,
1886–1988]

Araucaria hunsteinii
occurs in the highlands
of Papua New Guinea.
Most stands are in the
Morobe Province, with
some in the Central and
Eastern Highlands Prov-
inces. A few localities are
on the coast, but most
are well inland, mainly
in the Bismarck and Owen Stanley Ranges, with concentrations in the Bulolo and Wau Valleys
and their side valleys. Locality data based on herbarium collections seen by us are insufficient
to present an adequate map of the distribution of this species; the main reason for this is that
herbarium specimens are awkward to collect and press. Araucaria hunsteinii is often a very large
tree that can reach 90 m tall and 2 m or more trunk diameter. These tall trees are canopy emer-
gents and tend to grow in groves, undoubtedly related to episodic disturbance events after which
they could regenerate in large numbers, quickly filling the open space. The tallest trees grow in
moist sites with high rainfall. The altitude range is 550–2100 m so it is a species of lower montane
forests. This species is also planted as a forestry tree in the region, which makes mapping of the
natural distribution from herbarium specimens difficult if it is not clearly stated whether the tree
grew in natural forest or in a plantation, or planted in a village.

Agathis

The genus Agathis has 17 species, eight of these occur in Malesia. Sabah and Sarawak (Malay-
sia) in Borneo have four species. Another centre of relative species diversity is New Caledonia
with four species and there are three species in a small area near Cairns, Queensland, Australia.
Agathis robusta occurs in Papua New Guinea and in Queensland, Australia and is treated in the
chapter Australasia (map AU-8 on p. 348). The genus Agathis has an almost exclusively tropical
distribution, with the exception of one outlying species in New Zealand, A. australis. It extends
from N Sumatera and Peninsular Malaysia in the west to North Island in New Zealand and Fiji
in the east. Its northernmost occurrence is on Calayan, one of the Babuyan Islands N of Luzon
in the Philippines, at 19° 16’ N. Whereas the distribution in Malesia is fairly continuous (given
the archipelagic nature of the region), more isolated occurrences are found on the E coast of
Australia and in the SW Pacific. In Australia there are two locations that lie some 1200 km apart.
In the Pacific Ocean islands are obviously a limiting factor and there are wide distances of ocean
between New Caledonia, New Zealand and Fiji. But the Malesian species are far removed from
those in the SW Pacific also because they are absent from the main Solomon Islands and Bou-
gainville, only occurring on the Santa Cruz Islands, in Vanuatu, in New Caledonia and in Fiji, and
far to the south in New Zealand.
332 malesia

Map MA-66. The distribution of the genus Agathis.

Where did the genus Agathis originate? The oldest unambiguous fossils assigned to this genus
are from Australia and date from the Eocene (Hill & Brodribb, 1999), around the time when
Australia had separated from Antarctica but had a much more southern position. Phylogenetic
relationships of extant species still need further research, but initial results (Knapp et al., 2007)
suggest that A. australis of New Zealand is a basal species in this clade. Both lines of evidence
indicate a Gondwanan origin of the genus, which may have been present in Antarctica as well
as in Australia. In the Eocene the climate was almost globally warm, with average temperatures
now occurring in North Island, New Zealand, prevailing at high latitudes in Antarctica and
southern Australia. With the separation and subsequent northward drift of Australia into the
desert latitudes these trees, with the rainforests of which they are part, were driven to the NE
coast and when Australia connected with Malesia the genus could spread into the archipelago.
This scenario does not explain distant occurrences in New Zealand, New Caledonia and Fiji.
Probably the genus is much older than the Eocene; we just lack fossils of older age that can be
assigned to Agathis without doubt. Both New Zealand and New Caledonia were connected to
eastern Gondwana in the Early Cretaceous. Vanuatu and Fiji were not, so a capacity of dispersal
across the sea, also in Malesia, must be taken into account to explain the present distribution
of Agathis.
 malesia 333

Map MA-67. Agathis orbicula [n=8,

m=8, h=6, 1936–1991]

Agathis orbicula is a rare species

of Borneo. It is also poorly recog-
nized as distinct by field workers
and consequently its distribution
remains incompletely known. De
Laubenfels (1988) presented a
map in Flora Malesiana with two
dots on either side of the border
between Sabah and Sarawak and
three dots in central Sarawak.
Our map, based on specimens in
K and L (other herbaria only have
a duplicate of the type collection
D. J. de Laubenfels P614 also pres-
ent in K and L) gives four dots in
the border area between the two
states, one in N-central Sarawak
but not where De Laubenfels
placed his dots, and two further
S, in Indonesian Borneo (Kalim-
antan). There is clearly an identification problem with this species, especially when only foliage
of youngish trees is available. This species occurs in ‘kerangas’ forest on sandy soils at altitudes
between 450 and 1050 m, so it is a lowland tree. It is listed as Endangered (EN) on the IUCN Red
List due to logging and deforestation which take no notice of this species and its rarity.

Map MA-68. Agathis borneensis [n=102, m=97, h=4, 1865–1998]

334 malesia

Agathis borneensis is widely distributed in Borneo and in Peninsular Malesia and occurs less fre-
quently on the mountainous W-side of Sumatera (Aceh, Barisan Range). It is found in lowland
to upland tropical rainforest as scattered emergent trees and in low lying ‘kerangas’ forest on
sandy or sometimes peaty soils, where it can form extensive pure stands. Dependent on habitat
and age, it can be a very large tree 50 m or more tall with a clear bole up to 20–30 m and a maxi-
mum of 3.5 m diameter. A wide crown emerges from the general canopy of angiosperms mixed
with podocarpaceous conifers, the latter especially in ‘kerangas’ forest. The altitudinal range of
A. borneensis is substantial, from near sea level to ca. 2400 m, but with greater abuncance below
ca. 1200 m a.s.l. This species has been very heavily over-exploited in many areas and as a result
its total area of occupancy (AOO) is estimated to have at least been reduced by half and this is
still ongoing. Stands covering an estimated total of 30,000 ha discovered in Kalimantan in the
1930s had effectively been logged out by the mid 1960s. It is likely that many points on our map
representing older herbarium collections no longer represent existing stands of this tree. Agathis
borneensis is listed as Endangered (EN) on the IUCN Red List of Threatened Species despite its
wide distribution because of extensive logging.

Map MA-69. Agathis dammara [n=80,

m=77, h=18, 1741–2002]

Agathis dammara occurs in the

Philippines, Sulawesi and the
Moluccas. The two species A.
borneensis and A. dammara as
understood here and in A Hand-
book of the World’s Conifers (Far-
jon, 2010a) are not only separable
taxonomically (see textbox) but
appear to occupy distinct ranges.
Agathis dammara occurs to the
east of A. borneensis and there is
no sympatry. Wallace’s Line seems
to separate them between Borneo
and Sulawesi, but the Philippines
to the N are ‘west’ of Wallace’s
Line. The occurence of A. dam-
mara in the Philippines is perhaps
the result of more recent dispersal.
The habitat of A. dammara is simi-
lar to that of A. borneensis and it
can be a very large tree emerging
above the general canopy domi-
nated by Dipterocarpaceae or Fagaceae but usually mixed with numerous other trees. Exploitation
both of timber and of resin has depleted populations in many areas, especially in the Philippines. In
this country a ban on logging of ‘kauri’ has been imposed, but illegal logging goes on. Its presence in
forests still largely unexploited in the Moluccas and parts of Sulawesi render the conservation situ-
ation less severe than with A. borneensis. Whether this will last is a good question and the answer
depends on how well the Indonesian government can protect its remaining forests. Agathis dam-
mara is listed as Vulnerable (VU) on the IUCN Red List due to extensive logging.
 malesia 335

Agathis borneensis and A. dammara have been much confused in the literature as well as in the
field. Certain identification requires mature pollen cones; often these can be found on the forest
floor under the tree, but few will know that it is the character states of these little organs that
indicate the difference between the two species and not leaf size or shape, or the bark on the
trunk. In the herbarium, many specimens are sterile and cannot be identified with certainty.
However, when pollen cones were available, no specimen from Borneo could be assigned to
A. dammara. Because the other three species in Borneo occupy habitats in restricted areas, two
of them well known and collected, by extrapolation the common and widespread species can
be assumed to be A. borneensis, and this is what has been done for the Atlas. Agathis dammara
is the earliest known species; it was described from the Moluccas. It now includes A. celebica
and A. philippinensis, as was already understood by Whitmore (1980) but not accepted by De
Laubenfels in Flora Malesiana 10 (3) (1988). The pollen cones match those of A. dammara while
the leaf distinctions turned out to be spurious (Farjon, 2010a). Whitmore (1980) included some
montane populations in Borneo and Peninsular Malaysia with A. dammara, but these are now
recognized as neither belonging to A. borneensis nor to A. dammara.

Map MA-70. Agathis kina-

baluensis (black) [n=17, m=17,
h=5, 1915–1995]; A. flavescens
(red) [n=13, m=13, h=3, 1905–
1973]

These two species have

very limited distributions
and can be considered
rare. Agathis kinabaluen-
sis was based on a collec-
tion from Mt. Kinabalu
(D. J. de Laubenfels P625,
12 Aug 1978, A, K, L-holo-
type) and has been col-
lected on this mountain
in several places. However, it is not endemic there because it was later recognized to occur in
N Sarawak (4th Division) on Gunung Murud, from where it has been collected several times
between 1967 and 1995, and in the Kelabit Highlands around the village Pa Dali (H. M. Chris-
tensen 128, AAU, collected in 1992). Agathis kinabaluensis occurs in montane to upper montane
forest or scrub, on ultramafic substrates or on granite or sandstone. It is listed as Endangered
(EN) on the IUCN Red List. Agathis flavescens occurs on two mountains in Peninsular Malaysia,
Gunung Tahan and a short distance to the NW, Gunung Rabong. The altitude range is 1100–1900
m and the habitat ‘mossy’ forest or scrub. On Gunung Tahan, the highest mountain at 2189 m, the
species does not reach the summit, which is mostly covered in heath-like dwarf shrubs. Agathis
borneensis occupies a lower zone on this mountain covered with taller forest; the two species
are separated by a belt of vegetation devoid of Agathis. Gunung Tahan is in a national park and
remote, yet the species is considered at risk and listed as Vulnerable (VU) on the IIUCN Red List
mainly due to its limited occurrence, where a large forest fire could destroy one of the two known
populations.
336 malesia

Map MA-71. Agathis lenticula [n=19, m=19,

h=8, 1949–2001]

Agathis lenticula has a limited distribu-

tion, mainly in the mountains of Sabah
(Mt. Kinabalu, Crocker Range, Mt. Trus
Madi, Maliau Basin, upper drainage of
Padas River) and one locality, Gunung
Murud, in N Sarawak. In Flora Male-
siana (De Laubenfels, 1988) two years
after its formal description and naming
by David de Laubenfels, this species was
only known from Mt. Kinabalu and the
Crocker Range. More localities may yet
be found; as with other species in this
genus, the difficulty is partly in recog-
nizing this species as distinct from the
common A. borneensis in the absence of
pollen cones. Most ‘new’ discoveries were
made in the herbaria from specimens collected before the species was described in 1986 and only
one collection in the Conifer Database dates from after that year. We obviously need more botani-
cal collecting activity, bringing back specimens that can be identified, not just some twigs with a
few leaves. Agathis lenticula is an emergent tree in lower montane evergreen tropical rainforest. It
occurs in dipterocarp rainforest and forest dominated by Fagaceae at altitudes between 1050 m and
1700 m on a variety of dark to light soils. This species is listed as Vulnerbable (VU) on the IUCN Red
List due to unsustainable exploitation outside protected areas.

Agathis lenticula in the Crocker Range, Sabah (Malaysia) Borneo. Left: emergent tree; right: bole of the same
tree. Photo credit Aljos Farjon.
 malesia 337

Map MA-72. Agathis labil-

lardierei [n=48, m=45, h=4,
1792–1992]

Agathis labillardierei is
endemic to New Guinea
and nearby islands. It
is distributed from the
Bird’s Head Peninsula
in the west to the upper
drainage of the Sepik
River in the east, includ-
ing the islands of Yapen
and Biak. It is also indicated on a map (Fig. 76) in Flora Malesiana 10 (3) (De Laubenfels, 1988) to
occur on Misoöl Island and on Waigeo Island W of the Bird’s Head Peninsula, but we have not
found herbarium specimens to produce dots on our map. Agathis labillardierei occurs in tropi-
cal rainforest from near sea level to at least 2000 m; it is most common between 200 m and 1350
m. It eventually grows to a 50–60 m tall emergent tree on favourable sites and like several other
species of Agathis, regenerates abundantly after largee scale forest disturbance. Nearly all of these
young trees are later outcompeted by angiosperms, but a few emerge above the canopy and, as
long-lived trees, can survive for the next opporrtunity to regenerate.

Cupressaceae

The Cupressaceae are poorly represented in Malesia, with just one species in a monospecific
genus, Papuacedrus. This genus, now virtually confined to New Guinea, is known from the Oligo-
cene to Early Miocene of Tasmania and (with some uncertainty) from the Eocene of Antarctica
(Hill & Brodribb, 1999). The genus Papuacedrus is closely related to Libocedrus of New Caledonia
and New Zealand and to Austrocedrus and Pilgerodendron of S Chile and S Argentina. Libocedrus
is also known from the fossil record in Australia, including New South Wales and Tasmania.
Clearly, this is a group of taxa of Gondwanan origin with fragmentary distribution in the present
but more widespread in the past.

Map MA-73. Papuacedrus
papuana [n=114, m=109, h=8,
1913–2009]

Papuacedrus papuana has

two varieties, var. arfaken-
sis and var. papuana. The
differences are mainly in
the leaf shapes, but these
seem to become obscure
in the small leaves of
adult foliage on older
trees. Although var. arfak-
ensis seems to be more
common in the Bird’s Head Peninsula of New Guinea and may be the only variety so far found
in the Moluccas, we present them here on a map of the species without distinction. Papuacedrus
papuana is distributed from the Moluccas to the eastern end of New Guinea. The main distri-
bution follows the great central mountain chain of New Guinea; isolated populations occur on
Bacan Island and Obi Island (Moluccas), the Bird’s Head Peninsula, in the Cycloop Mts. and on
the Huon Peninsula. Whereas this species appears to be concentrated in certain regions of the
338 malesia

central mountains, the isolation on the map in western New Guinea (Papua, Indonesia) is at least
partly due to under-collecting compared to eastern New Guinea (Papua New Guinea). Papuace-
drus papuana is present in different forest zones from montane tropical rainforest to subalpine
scrubland, at altitudes of (620–)900–3600(–3800) m. Its greatest abundance is in ‘mossy’ forest
with nearly perpetual cloudy and rainy weather, which is at the highest altitudes reduced to a
tall ‘heath’ surrounded by tussock grassland, swamp and stands of tree ferns. Lower down in the
montane evergreen rainforest this conifer is more occasional, but trees can attain 40–50 m in
undisturbed primary forest. Common conifer associates are Phyllocladus hypophyllus, Dacrydium
and Podocarpus; in some locations it grows with Araucaria cunninghamii.

Pinaceae

The only genus of this Northern Hemisphere family that occurs naturally in Malesia is Pinus.
There are two species, Pinus kesiya and P. merkusii. Pinus kesiya occurs in N Luzon in the Philip-
pines, but is much more widespread in mainland SE Asia and is therefore treated in the chapter
Mainland Asia and Japan. That leaves one species to be treated in this chapter.

Map MA-74. Pinus merkusii

[n=27, m=26, h=12, 1844–1993]

Pinus merkusii is here treated

in a narrow taxonomic cir-
cumscription which excludes
populations in Indochina
commonly known as this
species; these are classified
as P. latteri and are mapped
in the chapter Mainland Asia
and Japan. Pinus merkusii
occurs in Sumatera and in
the Philippines; it is the only
species of pine that naturally
crosses the Equator into the
Southern Hemisphere, with
an isolated population near
Mt. Koerintji in Sumatera at
1° 42’ S. Its greatest extent is
in Sumatera, where it is abundant in the mountains of Aceh and S to Tapanuli; beyond there with
localities in Prapat, Sipirok and Mt. Koerintji the pines become more scattered. In the Philippines
there are two populations, the largest one in NW Luzon and a smaller one in Mindoro. Pinus
merkusii occurs in the mountains and forms open forests or pine ‘sanannas’ where grass fires are
frequent. Its higherst elevation is around 2000 m but most stands are below 1300 m; because it
is also planted for resin tapping its natural range is sometimes obscure. This species is certainly
not native in Jawa where many herbarium collections were made. The Dutch in colonial times
already had a hand in distributing this European-looking conifer with its useful resin and wood
from Sumatera to islands where it was inconveniently absent. Its natural stands have been dimin-
ished by logging and the species is listed as Vulnerable (VU) on the IUCN Red List.
Australasia

Geography

Australasia is a region of Oceania comprising Australia, New Zealand, New Guinea and neighbour-
ing islands in the Pacific Ocean. Here we exclude New Guinea, as it is treated within the biogeo-
graphic region known as Malesia. Relevant to the distribution of conifers, Australasia therefore
includes Australia with Tasmania (8.84 million km²), New Zealand (0.27 million km²) and the fol-
lowing archipelagos of small islands and islands in the Southwest Pacific: New Caledonia, Vanuatu,
Solomon Islands, Santa Cruz Islands, Fiji, Norfolk Island, and Tonga. A few other island groups
in this region have no conifers. Tonga is the farthest archipelago into the Pacific where conifers
occur naturally. All the islands with conifers are either parts of continental plates or accreted
terrains, or have emerged on sunken lands (rises) that were once connected to continents. True
oceanic islands in the Pacific, i.e. volcanoes that rose above the ocean surface from mid-oceanic
plates, do not have conifers. The region divides naturally into the continent of Australia, mostly
a desert land mass, and the islands, of which only New Zealand is of large size, comprising two
large and a number of small islands. Other islands are small or at most of medium size (the larg-
est, New Caledonia, is 18,576 km²). In this chapter, we will give accounts of conifer biogeography
separately for Australia, New Zealand, New Caledonia and the other Pacific Islands.

Conifers in Australasia

Families Genera Species Species + infra Countries with Area of occupancy Endemic taxa
conifers in km²
5/8 26/70 119/615 122/794 7/11 42,342/9,115,000 111
Numbers after ‘/’ are global or continent totals.

Map AU-1. Distribution of conifer species in

Australasia [Tonga not shown, but see map
AU-115 on p. 420]

The distribution map of Australasia shows

clearly that conifer species are concen-
trated on islands, especially Tasmania in
Australia, New Zealand, New Caledonia
and Fiji (see also map AU-2 on p. 342).
In Australia, concentrations occur in the
margins of the continent in Queensland,
New South Wales, Victoria and SW West
340 australasia

Australia. The islands are all mountainous and receive high levels of rainfall with limited dry
seasons. They are situated in the tropics and in the temperate zone of the Southern Hemisphere
(Norfolk Island, New Zealand and Tasmania) with the coolest climate in Tasmania and Stewart
Island (47° S). Australia extends from the tropics of northern Queensland to cool temperate Tas-
mania, but it has a vast interior that is arid to semi-arid, in which only a single conifer, Callitris
columellaris has a scattered occurrence. Rainfall is the limiting factor for conifer occurrence here,
not temperature.
At the species level of taxonomy, the island archipelagos and Australia all have unique conifer
floras, i.e. with very few exceptions no species occur in more than one of the geographical units
defined here. At the generic level there are numerous shared distributions, but also some unique
occurrences. This indicates separation of these areas for long periods of geological history. Links
at species level mostly occur with Malesia, in particular with New Guinea, reflecting the more
recent proximity of the two regions.

Table 4. The distribution of 26 genera of conifers occurring in Australasia across 10 areas.

The genera are arranged by family in Table 4. The first three are in Araucariaceae. This family, in
particular Araucaria, had a more global distribution in the Mesozoic and its Australasian distribu-
tion can largely be seen as relict. No seeds could easily disperse across wide stretches of ocean,
making dispersal events unlikely as explanations for current patterns of distribution. Agathis has
winged seeds, so perhaps short distance “island hopping” may have been possible within archi-
pelagos. It is at any rate the more widely distributed genus in the family, here occurring in six
of the 10 areas, mostly on islands. Wollemia, increasingly recognized in the fossil record of the
region since living plants were discovered near Sydney, New South Wales in 1994, is an extreme
relict reduced to virtually one location. The Cupressaceae are represented by genera 4 to 9 and
occur in Australia, Tasmania, New Zealand and New Caledonia, with most genera (and species)
in Australia. This family is the only one to occur in both the Northern and Southern Hemispheres,
but at least among living genera no southern representatives cross the equator. Tasmania has
two genera, Athrotaxis and Diselma, not found elsewhere. The Podocarpaceae, with genera 10–24
(some researchers also include Phyllocladus in this family) is the most diverse and widespread
family in Australasia. However, most are found on the islands, with only coastal Queensland with
its strip of tropical rainforest giving mainland Australia two extra genera beyond Podocarpus.
 australasia 341

Phyllocladaceae and Taxaceae are represented with one genus each, respectively in Tasmania and
New Zealand, and in New Caledonia.
The genus Podocarpus is spread across Australasia and is only absent from the small and iso-
lated Norfolk Island. In the family, it is the only large genus (ca. 100 species) with a pantropical/
Southern Hemisphere distribution. Its specialized seed cones are dispersed by birds, giving spe-
cies the potential to cross stretches of unsuitable habitat like seas. However, several other genera
in this family have similar seed dispersal strategies and yet are more restricted (Dacrycarpus,
Dacrydium, Prumnopitys, Retrophyllum), or even limited to a single area in this region. Dispersal
capacity does not guarantee dispersal and other factors must have played a role in these distri-
bution patterns. One major factor is the geological history of Australasia. In the Mesozoic the
main landmass in the Southern Hemisphere was an amalgam of all the present continents into
one, Gondwana. By the Early Cretaceous this had begun to break up and towards the end of that
period, ca. 70 Ma, separate continents had formed. The parting of Australia and Antarctica was
the last major break-up, with Tasmania linking the two as late as the Eocene ca. 50 Ma. New
Zealand, New Caledonia, Fiji and Tonga were all much closer to Australia and have drifted away,
while the Coral Sea and the Tasman Sea largely consist of submerged continental shelf. Con-
nections were severed by widening expanses of ocean, isolating plant populations and thereby
enabling speciation. This has caused uniqueness at species level in these areas and eventually led
to distinctions recognized as genera. Finally, extinction may also have played a role in the pres-
ent distribution patterns of genera. This is suggested by the peculiar distribution of Phyllocladus,
present in Tasmania and New Zealand and outside the region in Malesia (map AU-64 on p. 384
Its absence in Australia’s mainland and New Caledonia is probably due to extinction. More on
this will be said under the genus and species later in this chapter.
Australia, Tasmania, New Zealand and New Caledonia all have endemic genera, but the archi-
pelagos furthest away from Australia have none. Diversity also decreases with this distance. New
Caledonia is most diverse with 13 genera, four of these are endemic; these are remarkable numbers
given its modest size. New Zealand follows with 10 genera and three endemics. Given its vastly
greater size, Australia with eight genera is impoverished, with an equal number in nearby and
small Tasmania, and we shall expand on this separately under Australia. Actinostrobus and Wol-
lemia are endemic to the Australian mainland, but Sundacarpus is widespread outside the region
in Malesia. Tasmania is exceptional in that of its eight genera five are endemic to the island,
despite its proximity to the Australian mainland. The endemism of Tasmania is particularly inter-
esting in the light of the fossil record there (Farjon, 2008, p. 173). Many genera now present in
other areas of Australasia, e.g. Acmopyle, Agathis, Araucaria, Dacrycarpus, Dacrydium, Libocedrus,
and beyond (especially South America) once occurred there. The link with South America prob-
ably went over Antarctica, once ice-free and forested. The links with the Australian mainland and
Australasian islands demonstrate that these genera were once more widespread in the region.
At the species level almost all taxa are endemic to Australasia, a mere 11 species are also dis-
tributed in Malesia; two, Agathis robusta and Araucaria cunninghamii with a different variety,
so at that level they are also endemic. The other species shared with Malesia are Dacrycarpus
imbricatus, Dacrydium beccarii, D. magnum, D. nidulum, D. xanthandrum, Podocarpus glaucus,
P. neriifolius, P. pilgeri, Retrophyllum vitiense and Sundacarpus amarus. All have very wide distri-
butions in Malesia and just happen to extend into Australasia (Queensland in the case of Sun-
dacarpus amarus and the Solomon Islands to Fiji for all other species). We can consider these to
be Malesian species that got an opportunity to spread into the region when islands or landmasses
had moved close enough. Few species seem to have made the journey in the other direction, if
we assume that the centre of origin lay within the larger area of occurrence, which on statistical
grounds would be the default assumption unless there is evidence to the contrary. The excep-
tion is Araucaria cunninghamii, related to its congeners in New Caledonia and Norfolk Island. It
spread to New Guinea and evolved to a slightly distinct taxon (variety), joining another species
probably already there which is related to Araucaria bidwillii in Queensland. The level of ende-
mism in Australasia is therefore extremely high (96% at species level) and not exceeded by any
342 australasia

other ‘continental’ region defined in this Atlas. Extreme and long isolation is the major cause of
this, and while Australia became impoverished due to desertification, in earlier times it was much
greener (White, 1994a, 1994b) and must have served as a major link to Gondwana, where many of
the genera in this region probably originated.

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Map AU-2. Map of Australasia showing numbers of conifer species per 0.5 degree cell. Conifer diversity is
highest on the islands, New Caledonia being the most diverse. In Australia, only the area around Cairns in
N Queensland and W Tasmania score higher than seven species per 0.5 degree cell.

Map AU-3. Rarity values per 0.5 degree cell. Cell score is the total of the rarity calculations for each species
in the cell. The rarity calculation is 1 divided by the total of cells occupied by a species. Thus, a species that
occurs in one cell scores 1 while a species that occurs in 10 cells scores only 0.1. The highest scoring cell in
Australia is in N Queensland, most narrow endemics are in New Caledonia.

The diversity analysis shown in map AU-2 demonstrates the relative poverty of the Australian
continent, where only a limited area (one cell) near Cairns in N Queensland has more than seven
species per cell. This is due to the occurrence of a few species of Agathis and some in Podocar-
paceae with limited ranges; hence this same grid cell scores high in the rarity values calculated
for map AU-3. Tasmania is slightly more diverse, with several cells containing 8 or more species,
all in the western highlands part of the island. Several of these species have limited ranges within
Tasmania, and therefore elevate the rarity score in map AU-3. On the mainland, two cells in New
South Wales score similar rarity values as Tasmania, due to Pherosphaera fitzgeraldii (Podocar-
paceae) and Wollemia nobilis (Araucariaceae) both with very limited distributions. New Zealand
scores higher than Tasmania on the species diversity scale of map AU-2, but its scores for rar-
ity fall below it because all species have relatively wide distributions. Endemism for conifers in
 australasia 343

New Zealand is at a national level, not a local level. By the same measures Norfolk Island with
a single species (Araucaria heterophylla) scores very low in conifer diversity but its rarity score
is high as this species occurs nowhere else naturally (it has been very widely planted). Species
diversity is easily highest in New Caledonia and the rarity score is at the highest values in many
of its 0.5 degree cells due to much local endemism. Finally, while diversity is limited on the other
islands in the SW Pacific, Fiji has moderately high scores for rarity.
Important conclusions follow from these two maps. One is that the islands of the SW Pacific
within the Australasian Region have disproportionate high levels of diversity and endemism for
conifers. The continent of Australia scores very low in comparison. Conifer biodiversity in Aus-
tralasia is concentrated on the islands and these islands are all isolated from major land masses.
Conservation of this biodiversity, which is in biogeographic and phylogenetic terms unique to the
region, has to be a high priority.
344 australasia

Australia

Geography

Australia (here including Tasmania) is an island continent situated entirely in the Southern
Hemisphere with an area of 8.84 million km². It is an ancient fragment of the supercontinent
Gondwana which became separated from Antarctica in the Late Cretaceous/early Tertiary before
it drifted northwards. This drift took it largely out of a cool temperate climate zone through an
arid zone and up to the tropical zone. Lack of major orogenetic activity has resulted in a relatively
low, eroded continent characterized by ‘inverted’ landscapes and many rivers ending in interior
evaporation basins. As a result, much of the interior of the continent has a desert or semi-desert
climate and significant rainfall is restricted to the margins, from the tropical Arnhem Land and
Queensland coast down to New South Wales, Victoria, coastal parts of South Australia and the
SW corner of Western Australia. Summer rains prevail in the north, winter rains in the south and,
most markedly, in the southwest. The highlands of SE New South Wales/Victoria and the island
of Tasmania have a more evenly distributed rainfall. The Great Dividing Range is situated along
the entire eastern margin of the continent, culminating in Mt. Kosciuzko (2229 m) in its south-
ernmost part. There are no other substantial mountain ranges with a marked influence on climate
and vegetation. Rainforest is present along the eastern Queensland coast, in fragments along the
coast of New South Wales, and in western Tasmania, ranging N-S from tropical to cool temperate.
Other forests and woodlands are dominated by Eucalyptus (Myrtaceae) and in the interior of the
continent also by Acacia (Leguminosae).

Conifers in Australia

Families Genera Species Species + infra Countries with Area of occupancy Endemic taxa
conifers in km²
4/8 14/70 41/615 41/794 1/1 21,325/8,840,000 40
Numbers after ‘/’ are global or continent totals.

Only four families are represented in Australia: Araucariaceae, Cupressaceae, Phyllocladaceae and
Podocarpaceae. Of Araucariaceae there are three species of Agathis, three species of Araucaria,
and Wollemia nobilis; of Cupressaceae the genus Actinostrobus has three species, Athrotaxis has
three species, Callitris is represented with 13 species, and Diselma archeri is monospecific; Phyl-
locladaceae is represented by Phyllocladus aspleniifolius; in Podocarpaceae occur seven species of
Podocarpus, two in Pherosphaera, Lagarostrobus franklinii, Microcachrys tetragona, Prumnopitys
ladei and Sundacarpus amarus. This sums up the native conifers of Australia, including remote
Norfolk Island with Araucaria heterophylla. Of these, the genera Athrotaxis, Diselma, Lagarostro-
bos, Microcachrys and Phyllocladus aspleniifolius are endemic to Tasmania. The only genus that
is widespread in Australia is Callitris, of which most species are well adapted to seasonal aridity,
heat and above all, fire. This adaptation has enabled its spread and adjustment to specific niches
and localities, leading to speciation.
It is obvious that conifers are poorly represented in Australia, comparable to Africa in numbers
of taxa present. The level of endemism is very high (97.5%) with only one of Australia’s conifer
taxa (Sundacarpus amarus) occurring also in Malesia. Two other conifer species, Araucaria cun-
ninghamii and Agathis robusta, have sister varieties in New Guinea. Were it not for Tasmania,
Australia would be as poor in conifers as Africa. This is reflected in Table 4, where for this reason
we treated Tasmania as a separate area, leaving mainland Australia with only eight genera. We
must now discuss where they occur, and why.
 australasia 345

Map AU-4. The distribution of coni-

fer species in Australia.

The overall pattern of distribution

is characterized by concentrations
around the margins of Australia,
with scattered occurrence else-
where and large empty areas. In
Queensland, conifers are abun-
dant on the Atherton Tablelands
and other upland areas around
Cairns, as well as in the SE corner
of the state in mountains and
tablelands to the east of Brisbane.
In New South Wales, conifers have
a more scattered distribution, thin-
ning out to the west across the
Great Dividing Range. Another, but more diffuse, concentration of conifer distribution is found
on either side of the border between Victoria and South Australia. In Tasmania most conifer spe-
cies occur in the central and western highlands. A wide gap, in the south occupied by the Nul-
larbor Plain, separates eastern from western conifer distribution. In Western Australia conifers
occur in several areas of the SW corner of the state. In the tropical north, the interior and the NW
coast of the continent, conifers are very scattered or absent altogether.
This distribution of the conifers of Australia fits very well with a map showing the main rainfall
patterns of the continent. The empty areas and those with only a scattered occurrence of conifers
coincide perfectly with the arid zone and with the semi-arid zone with summer rainfall. Only
coastal and upland areas with tropical rainforest in the moist tropical zone have concentrations
of conifer species, these occur on the NE coast of Queensland. Elsewhere, conifers are found in
moist temperate areas with hot or warm summers, which have mostly uniform rainfall through
the year, and in areas with winter rainfall.

Map AU-5. Map of Australia show-

ing seasonal rainfall patterns. Source:
Australian National Herbarium
CANB; Government of Australia.

It appears therefore, that climate

is the principal component of the
present environment explaining
much of the general distribution
pattern of conifer species in Aus-
tralia. Soil conditions, related to
geology and climate, are another
limiting factor. In Australia,
widespread deep weathering has
chemically altered sediments
during wetter periods in the
Palaeocene to Eocene, 65 to 35 Ma, when Australia was green and forested (White, 1994b). Com-
plex processes of mineral movements and concentrations in the profile have resulted in ‘duricrust’,
iron-based (ferricrete) or silica-based (silcrete) hard layers. After erosion of softer strata mainly dur-
ing drier times, these layers are now forming caps on top of mesa-like plateaus, creating ‘inverted’
346 australasia

landscapes, i.e. land that once formed flat depressions and shallow valleys is now the most elevated
part of the terrain. These duricrust areas are extensive and not limited to the present arid zone; they
are widespread also in the moist tropical zone. This northern part of the continent has arrived in
the tropics geologically recently, after passing through the arid zone, due to Australia’s rapid drift
northward after it finally broke away from Antarctica. These duricrust soils are difficult or unfit for
plant growth; few species have been able to adapt to this; among conifers one or two species of Cal-
litris can sometimes be found on them, often with a severely stunted growth habit. Deep weathering
also creates horizons above the water table from which all minerals have leached out, creating often
chalk-white and fine-grained sediment. Erosion has spread this unfertile material from ‘breakaways’
below the duricrust caps across the now lower surrounding land, creating more terrain unfit for
prolific vegetation even when rainfall might have been sufficient. Australia’s major rivers, due to
the topography of the continent with raised margins on several sides, are inward draining. Many do
not reach the oceans and their water evaporates leaving extensive salt playas, the largest of which
is Lake Eyre in South Australia. Salt playas are also common in the SW corner of Western Australia,
where winter rainfall allows vegetation to develop, including conifers such as Actinostrobus and
Callitris. Salt pans and playas are expanding there, partly due to clearing of vegetation for wheat
growing and possibly also due to climate change. Conifers generally do not tolerate salt. Finally,
Australian vegetation for the most part is highly flammable and burns frequently. Eucalypts, wattles
(Acacia) Banksias and other members of Proteaceae are all adapted to this in various ways and
dominate the vegetated landscape. Few conifers have evolved adaptations to fire, most prominently
pines (Pinus, not native) and members of Cupressaceae. Foremost among the latter in Australia is
the genus Callitris and this is indeed the most widespread and diverse genus in the continent. Acti-
nostrobus from Western Australia has a similar tolerance; the most common strategy is to preserve
the seeds in fire resistant closed cones. Almost a peculiarity among Podocarpaceae, Podocarpus
drouynianus in SW Western Australia is a low, multi-stemmed shrub that resprouts from the base
after fire, a tactic adopted by most eucalypts and wattles. All other conifer species must avoid fires
and this is a further explanation for their retreat to moister coastal regions.
It is appropriate to speak of a retreat. There is abundant fossil evidence of conifers as a promi-
nent component of the Australian flora in past geological periods (White, 1994a; Hill & Brodribb,
1999). This fossil record, from the Jurassic to the Pliocene, is evidence for conifers, some extinct,
others extant but ‘marginalized’ to Tasmania or coastal Queensland, to have occurred in places
where the climate no longer suits them. In Australia, conifers have been fighting a rearguard
battle against climate change and its associated hazards. The arrival of people on the continent
resulted in an increase of fires and while this has been ongoing for tens of thousands of years, the
vegetation changed dramatically in favour of grasslands and savannas with eucalypts against for-
ests, as pollen records show. Finally, the late arrival of Europeans added logging for timber and
clearing of forest for agriculture.

Callitris columellaris, a solitary tree hold-

ing out in the desert landscape of Austra-
lia’s arid centre (Kings Canyon National
Park). Its glaucous foliage is probably an
adaptation to intense solar radiation.
Photo credit Aljos Farjon.
 australasia 347

Agathis

Agathis is one of three genera in the family Araucariaceae, all are present in Australia. This genus
is almost entirely tropical, with only one species, A. australis, outside the tropics in New Zealand.
Its main distribution is in Malesia and the SW Pacific (for a map of the genus, see map MA-66
on p. 332), but three species occur in near coastal areas of Queensland in Australia. Almost all of
the 17 species grow to very large trees and are usually emergents, rising their crowns eventually
well above the forest canopy. The three Australian species are in this category. In the past, all
three were heavily exploited for their valuable timber, but this exploitation has almost ceased. It
is unclear to what extent this has led to reductions in their ranges, but it has certainly reduced
numbers of large trees and may have removed almost all of the forest giants. Large trees still
exist, especially of A. microstachya, but it will take a few centuries for most other trees to attain
full size.
Agathis microstachya has the smallest range of the three Australian species. It is confined to the
Cook District, occurring in an area S and SW of Cairns, ranging from the heights of the Atherton
Tablelands at 1100 m towards but not reaching the coast at 400 m altitude. Large trees can be seen
around Lake Barrine, largest of the two volcanic crater lakes in Crater Lakes National Park. This
species appears to be associated with soils derived from volcanic rock, which are abundant in the
area, but limit its range. High annual rainfall, up to 3000 mm, can be another limiting factor. After
logging was abandoned, an estimate gave the total number of mature trees as fewer than 10,000,
but most of these are now within protected areas and the decline has ceased.

Map AU-6. Agathis microstachya [n=16, m=14, h=2,

1915–1995]

Agathis microstachya near Lake Barrine, N Queensland. The tree fills a gap in tropical notophyll vine forest,
characterized by lianas (‘vines’) which connect trees. If a large tree falls, it pulls down others, and in such
a large gap Agathis can grow up fast, eventually emerging above the canopy and spreading its wide crown
to fill the space. Photo credit Aljos Farjon.
348 australasia

Map AU-7. Agathis atropurpurea [n=10, m=10, h=2,

1923–1988]

Agathis atropurpurea occurs in the same area

of Cook District as A. microstachya, but also
on Mts. Misery and Spurgeon NW of Cairns. It
does not grow on volcanic (basaltic) rock but
on granite, which forms domes that rise above
the volcanic formations, giving A. atropurpurea
an altitude range of 900–1500 m. This species is
rare and individual trees in the forest can occur
far apart. It is recognizable by its distinct purple
bark, exposed when older grayish flakes peel off.
Wooroonooran (Wurunuru) National Park is a particular stronghold for this species.

Map AU-8. Agathis robusta ssp. robusta (black) [n=23,

m=23, h=7, 1859–2003]; Agathis robusta ssp. nesophila (red)
[n=16, m=15, h=4, 1956–1982]

Agathis robusta has two subspecies, ssp. robusta in

Queensland, Australia and in the Port Moresby area
of Papua New Guinea, and ssp. nesophila in the high-
lands of Png and on the island of New Britain. The
distribution in Queensland shows a familiar disjunc-
tion [see Araucaria bidwillii (map AU-9 on p. 349)
and Callitris macleayana (map AU-26 on p. 359)], with
populations near Cairns in the north and near Bris-
bane in the south, separated by ca. 1100 km. The third
population of ssp. robusta near Port Moresby is 900
km N of Cairns. Such disjunctions in a species with
no effective mechanisms for long distance seed dis-
persal almost certainly indicate relict occurrences. Is
there a difference between the two coastal locations
in Queensland? The northern area is tropical, but the
southern area is (warm) temperate and rainfall pat-
terns differ. Port Moresby is only 9° 30’ S of the Equa-
tor and the population near it experiences a similar
climate to that near Cairns. If relict, we should accept
that the remaining three disjunct populations in Aus-
tralia and New Guinea are accidents of history, in which extinction played a major role.
Agathis robusta ssp. robusta, in its southernmost location, occurs in semi-evergreen forest from
near sea level on Fraser Island to ca. 900 m, where occasional frost is recorded. In N Queensland
it occurs in notophyll vine forest, characterized by evergreen trees with small leaves and numer-
ous lianas. Here it is partly sympatric with A. microstachya but occurs at lower altitudes down to
near sea level. In Png, in the vicinity of Sirinumu Lake, a large reservoir supplying electricity to
Port Moresby, it is in mixed tropical low montane forest (600–850 m) and has been selectively
logged for some time.
Agathis robusta ssp. nesophila is found in lower montane tropical forest in Png in the Eastern
Highlands, the Owen Stanley Range and on the large island of New Britain. Its altitudinal range is
greater than that of ssp. robusta, between 400 m and 1900 m. It grows on a variety of nutrient-poor
soils derived from granite or other igneous rock, including ultrabasic types.
 australasia 349

Araucaria

The genus Araucaria is represented by two species in the continent of Australia, one of these,
A. bidwillii, is endemic to Queensland, the other species, A. cunninghamii has two varieties,
A. cunninghamii var. cunninghamii in Australia and var. papuana distributed widely in New
Guinea. We will treat this species and its varieties here under Australia. The third Australian spe-
cies is A. heterophylla, in the wild only occurring on Norfolk Island, which is Australian territory
but lies 1400 km E of the continent in the Tasman Sea. We shall deal with it under “interarc
islands” later in this chapter. Araucaria has a disparate distribution in South America, Australia,
New Guinea and New Caledonia (for a map of the genus see map AU-74 on p. 391), but a long
fossil record shows that the genus has been present on most continents. The two Australian spe-
cies treated here are good examples of relict species, in particular A. bidwillii which has been
classified in its own section Bunya. Further reduction of ranges and abundance were caused more
recently by logging, but the remaining stands of these trees are now mostly in protected areas.

Map AU-9. Araucaria bidwillii [n=25, m=21, h=11, 1843–

2002]

Araucaria bidwillii is an iconic tree in Australia; it was

a major food tree for Aboriginal tribes who held large
gatherings in good crop years to collect the seeds from
the massive cones. Its distribution in Queensland is dis-
junct, with the major populations in the SE of the state
and only a few scattered trees at Cannabullen Falls and
Mt. Lewis in N Queensland, ca. 1000 km distant. In the
SE it occurs in five main areas: the Blackall Ranges W
of Nambour; the Bunya Mountains W of Yarraman; the
upper reaches of the Brisbane River; the upper Mary
River Valley and the Yarraman/Blackbutt area. There
are slight differences between the northern and south-
ern populations, but these are probably not taxonomi-
cally significant. This species, known as the Bunya pine,
is a magnificent tree to 45 m tall, with a straight bole to
1.75 m in diameter. Its wide spreading umbrella-shaped
crown overtops surrounding angiosperm trees, making
the conifer an emergent in the notophyll vine forest.
It can also be gregarious with A. cunninghamii, which
is potentially taller but with a narrow crown. In the
Bunya Mountains, today its major stronghold, balds, i.e. areas of coarse grassland occur, sharply
defined from the surrounding forest. Slowly, A. bidwillii is seen to encroach on these balds, which
could mean that they were maintained by fires lighted by people.
350 australasia

Trees of Araucaria bidwillii raise their

broad crowns above the canopy of
the forest in the Bunya Mountains
National Park, SE Queensland. While
seedlings do occur in the forest, few
seem to be succesful and regenera-
tion depends on the creation of open
spaces such as the ‘bald’ seen in the
foregound. Photo credit Aljos Farjon.

Araucaria cunninghamiii in Australia has a more or less continuous distribution from near Grafton
in NSW north to near Townsville in Queensland, in its southern part more widespread but increas-
ingly following the coast northward, including on islands, and becoming rare N of Cairns. This
variety occurs at low altitudes from near sea level to around 900 m. It can occur as solitary trees
in dry vine forest in coastal areas on white sand, but also form quite extensive stands in moun-
tains further inland, e.g. in the McPherson Range along the border between Queensland and NSW
Araucaria cunninghamii has wind-dispersed seeds and can colonize open areas, e.g. after fire,
effectively. In a later phase of succession, other trees may replace it save for some large trees that
are emergent. In New Guinea A. cunninghamii var. papuana occurs across the entire island, from
the D’Entrecasteaux Islands to the Birds Head Peninsula, but is absent in the southern lowlands.
Its altitudinal range is extensive, from ca. 200 m in some localities near the N coast to 3355 m
(the highest herbarium record) in the central mountains. It is most common in montane forests
dominated by Castanopsis, Lithocarpus, Cinnamonum, Calophyllum and Schizomeria.

Araucaria cunninghamii var. cunninghamii on a ridgetop in

Queensland. Eucalyptus forest covers the lower slopes and
grassland, resulting from fires, replaces the forest below.
On the ridge, poorer soils and higher rainfall favour the
conifers. Photo credit Aljos Farjon.

Map AU-10. Araucaria cunninghamii var.

cunninghamii (black) [n=44, m=34, h=12,
1802–2000]; A. cunninghamii var. papuana
(red) [n=83, m=67, h=10, 1909–1995]
 australasia 351

Wollemia

Wollemia nobilis is the single species in the recently (1994) discovered genus Wollemia, named
after its location, Wollemi National Park in NSW, NW of Sydney. It is restricted to an extremely
small area, only a few km² in extent, but its exact locality is not publicly known, so the dot on
the map is only an indication of its whereabouts. Its habitat is a deep sandstone canyon, with its
tributary gorges, at the narrow bottom of which are perennial streams allowing a strip of dense
forest dominated by Ceratopetalum apetalum and Doryphora sassafras, two trees in the families
Cunoniaceae and Monimiaceae. Ferns dominate the ground flora.

Map AU-11. Wollemia nobilis. [n=1, m=1, h=2, 1994] Here shown in its natural environment in a canyon in
Wollemi National Park, NSW. New trunks are continuously reiterated from the base of old trees, forming
clusters of stems that are connected to the parent tree. Not leaves, but entire branches are shed, in the
upper crown after male or female cones have been produced at their tips. A moist microclimate and steep
canyon walls protect the trees from fires that may rage above in the eucalypt woodland. Photo credit
J. Plaza.

There are three tiny subpopulations of W. nobilis a short distance apart; within these the trees
appear to be regenerating from their base and seedlings, although present, are rare. These can-
yon bottoms are protected from fires, which frequently occur on the upper canyon sides and
the plateau on top, where the forest is dominated by eucalypts. The Wollemia trees are tall and
eventually emergent. This and the coppicing capacity indicate a positive feedback with periodic
fires, without which the angiosperm trees would take over. However, the three subpopulations
are now so small that it is uncertain whether this ecosystem still functions as it may have in the
past. Wollemia nobilis is listed as Critically Endangered (CR) on the IUCN Red List. Fossil evi-
dence, recognized in pollen deposits and also macro-fossils since its discovery, indicates a much
wider past distribution, perhaps as far as New Zealand during the Late Cretaceous (Kunzmann,
2007). The chance survival of Wollemia in a gorge surrounded by an environment hostile to it
must count as one of the remarkable events in the history of life.
352 australasia

Actinostrobus

Map AU-12. The distribution of the genus Actinostrobus in

Australia.

The genus Actinostrobus (Cupressaceae) is endemic to

Western Australia and comprises three species. It is
morphologically distinct from Callitris, but morpho-
logical and anatomical evidence used in cladistic anal-
ysis indicates that it may be derived from within the
Callitris clade (Piggin & Bruhl, 2010). Under cladistic
principles the genus cannot be maintained as it makes
Callitris ‘paraphyletic’, but from an evolutionary point
of view the hypothesis that the ancestor of Actinostro-
bus belonged to Callitris does not invalidate it as a
genus. We deviate from the insistence on ‘monophyly’
in this case and maintain Actinostrobus as a valid
genus. Its species are distributed in a limited near-
coastal region of SW Western Australia.

Map AU-13. Actinostrobus pyramidalis [n=37, m=25, h=11,

18??–1997]

Actinostrobus pyramidalis is an erect, often pyramidal

shrub to 6 m tall, often growing in gregarious groups.
It is locally common in low, open dwarf scrub (‘kwon-
gan’), or in low woodland dominated by Eucalyptus
spp. (‘mallee’); often in road verges through farmland
in disturbed vegetation. It tolerates poor soils and
even light levels of salinity. It occupies areas now
under increasing urbanisation, which leads to losses
of subpopulations. On the other hand it may also
spread along roads to new areas.
 australasia 353

Map AU-14. Actinostrobus arenarius Actinostrobus arenarius near Geraldton, Western Australia.

[n=30, m=25, h=10, 1901–1997] Photocredit Aljos Farjon.

Actinostrobus arenarius is the most widespread of the three species in this genus, but does not
occur as far south as A. pyramidalis. It is found from Lake Grace north to the Murchison River, a
distance of 600 km. In the country between Perth and Geralton it is most common, occurring in
low, open dwarf scrub (‘kwongan’), often on road verges where broad strips of the native vegeta-
tion are preserved. It is readily colonizing disturbed ground along roads and in abandoned fields,
often forming extensive stands of upright, columnar shrubs.

Map AU-15. Actinostrobus acuminatus [n=20, m=15, h=11,

1859–1997]

Actinostrobus acuminatus is a small, often creeping

shrub usually half concealed under blown sand, but
sometimes erect to 4 m tall. It is found in low scrubland
(‘kwongan’) on plains in dry or moist leached white or
grey sand or clay usually over laterite (hardpan). It is
also found in taller vegetation dominated by eucalypts
(‘mallee’) where it can be more erect. It is most com-
mon in the sandy plains between Three Springs in the
north and Dandaragan in the south, with abundant
plants in Lesueur National Park, a reserve protecting
the ‘kwongan’ vegetation, comparable in many ways
to the ‘fynbos’ of South Africa. Fires burn regularly in
this vegetation and the half-buried shrubs of A. acumi-
natus are probably well protected here.

Callitris

The genus Callitris (Cupressaceae) occurs mainly in Australia, but two of the 15 species are endemic
in New Caledonia. In Australia, it is by far the most diverse and widespread genus of conifer, with
most species having evolved adaptations to the climatic and edaphic conditions, including fire,
that prevail in Australia as described in the introduction to the continent (pp. 344–346). One
species, however (C. macleayana) occurs in wet sclerophyll forest and subtropical rainforest on
the east coast.
354 australasia

Map AU-16. The distribution of the

genus Callitris.

The distribution of Callitris gener-

ally compares well with the distri-
bution of all conifers in Australia.
Differences are apparent in Tas-
mania, where most conifers are in
the centre and west of the island
but Callitris, adapted to drier con-
ditions, prevails in the east. In
Queensland, while Callitris occurs
in these areas, the concentrations
of conifers near Perth and Brisbane seen on the all-species map of Australia (map AU-4 on p. 345)
are no longer present here. In all other states, Callitris either strongly dominates (Western Aus-
tralia, New South Wales) or entirely makes up the distribution of conifers in the continent. This
genus can be considered successful in having spread widely to occupy suitable habitat, diversify-
ing into 13 species. A number of these have themselves wide ranges, but others are restricted to
small areas and in some cases to rare local habitats. The growth habit of different species varies
from dense shrubs to tall trees; this is generally congruent with the type of vegetation in which
the species occur. Forests in eastern Australia along the Great Dividing Range and on the coast
have mostly tree forming species; ‘mallee’ and scrub harbour low, multi-stemmed taxa. Extreme
conditions of soil, as prevail on granite outcrops and on ‘breakaways’ or sandstone plateaux
can be the habitat of dwarfed forms like Callitris muelleri and C. monticola and these are often
very local species. Other species have wider ecological tolerance or are even opportunistic invad-
ers of disturbed vegetation or soil, as in pasture land or in road cuts. Their distributions cover
large areas and are perhaps even expanding into areas where they were absent before European
settlement.

Map AU-17. Callitris columellaris

[n=175, m=160, h=17, 18??–2006]

Callitris columellaris is the most

widespread species in the genus.
The taxonomy here accepted
includes C. glaucophylla and C.
intratropica (Farjon, 2005a) which
were treated as separate species in
Flora of Australia 48 (K. D. Hill in
McCarthy, ed. 1998). On the coast
of Queensland and New South
Wales between Bribie Island and
the Bundjalung National Park (27°
03’ S to 29° 20’ S) it occurs mostly
on or behind coastal sand dunes.
This is C. columellaris sensu Flora
of Australia. In the interior of these states and further west foliage colour often changes from
green (near-coastal populations) to glaucous, with very glaucous forms found e.g. in the MacDon-
nell Ranges and other localities in the centre of Australia (see photo p. 346). This phenomenon
can be found in other species of conifer and is associated with high levels of solar radiation (UV
light), either in deserts or at high altitude. Individual trees in a population can vary much in this
aspect. Callitris columellaris occurs on a wide range of substrates but all are characterized by a
 australasia 355

deficiency of nutrients; this species, probably better than most in its genus, can survive under
extreme conditions of climate and soil.

Callitris columellaris near Quirindi, New South
Wales forms open woods with grasses on the
ground, maintained by frequent light fires and/or
grazing of livestock. Photo credit Aljos Farjon.
Callitris columellaris near Scone, New South Wales.
Here trees with fastigiate branching, forming a
columnar habit (photo) appeared to occur side-
by-side with broad-crowned trees as seen in the
previous picture. In the background is eucalypt
woodland. Photo credit Aljos Farjon.

Map AU-18. Callitris preissii [n=169,

m=155, h=14, 18??–1997]

Callitris preissii is here also

treated in a wider sense than in
Flora of Australia 48 (K. D. Hill in
McCarthy, ed. 1998) and includes
C. gracilis and C. tuberculata (Far-
jon, 2005a). Like C. columellaris its
distribution spans the continent
from west to east, but it is with
a few exceptions limited to the
areas with winter rainfall. A wide
gap of about 1200 km separates
western from eastern popula-
tions; this area is mostly occupied
by the Nullarbor Plain where no
conifers occur. This plain was
once part of the submerged continental shelf bordering the Southern Ocean and was elevated to
a few tens of metres above sea level to become an extremely flat limestone area 200,000 km² in
extent. Harsh conditions only allow a sparse dwarf scrub vegetation with clumps of spiny grasses
in between. On sandy shores and islands near the coast in SW Western Australia C. preissii devel-
ops into a tree, but not of great stature and frequently multi-stemmed from a short trunk. Away
from the coast on lateritic soils a shrub form prevails (C. tuberculata) but on inland sand plains
and dunes tree forms appear again. The largest trees are found in such dune landscapes in NE
Victoria. Dunes often bury accumulated litter which on slow decomposition provides nitrogen
to plants, often resulting in more luxurious growth than is possible in the leached or ‘duricrust’
soils surrounding them.
356 australasia

Map AU-19. Callitris verrucosa [n=66, m=62,

h=13, 18??–1997]

Similarly distributed as C. preissii but not

reaching the coast in Western Australia
is C. verrucosa. This species is often con-
fused with C. preisii on herbarium labels,
so only specimens seen are mapped
here. It is most common in the sand
dune areas south of and near the mouth
of the Murray River and in the Little
Desert of Victoria. Here it can be grow-
ing together with C. preissii, with that
species best developed on the dunes and
C. verrucosa commonly on the flat
ground between them. As with the previous species, there is a wide tolerance of substrate diver-
sity and adversity. Callitris verrucosa if it grows to tree habit remains smaller than C. preissii even
where growing in the same area.

Map AU-20. Callitris canescens [n=47, m=42, h=8, 1830–

2011]

While more restricted in its range than the previous

two species, C. canescens also has two subpopulations
west and east of the Nullarbor Plain. It is most abun-
dant in the semi-arid winter rain zone of Western
Australia. Growing in dwarf scrub (‘kwongan’) and
Eucalyptus shrubland (‘mallee’) it remains of low stat-
ure and usually a shrub. Both vegetation types are rich
floristically, with numerous species of the families
Myrtaceae and Proteaceae and the well-known ‘grass
trees’ (Xanthorrhoea) all adapted to fire. These veg-
etations have been extensively destroyed (grubbed)
to make way for wheat growing and other uses, but
often broad strips along roads were spared. As a result
this species is still widespread even though reduced
in numbers.
Callitris canescens in roadside ‘mallee’ vege-
tation, Western Australia. Photo credit Aljos Only two species of Callitris are limited to Western
Farjon.
Australia, C. roei and C. drummondii. They have similar
habitats, i.e. ‘mallee’, ‘Melaleuca thicket’ and occasionally ‘kwongan’ as well as disturbed ground
alongside roads, where they may invade, or margins of salt lakes and flats above the periodic
level of water after rains. Yet their distributions only partly overlap, with C. roei distributed more
inland and C. drummondii limited to a broad coastal strip along the Southern Ocean from Albany
in the west to Cape Arid in the east. Both are distributed within the zone of winter rain, but
annual precipitation falls sharply from 1400 mm near Northcliffe on the SW coast to only 300 mm
in the ‘wheatbelt’ which stretches roughly east of Perth into the interior of SW Western Australia.
Callitris roei still occurs there, as well as in some localities on the south coast, but C. drummondii
is absent. It is therefore likely that the two species differ in their tolerance of drought, which is
more severe inland with less water available from winter rains. Callitris roei is listed as Endan-
gered (EN) on the IUCN Red List, mainly due to large-scale conversion of natural vegetation in
the ‘wheatbelt’ and deterioration of remaining habitat with rising levels of salt in the soil.
 australasia 357

Map AU-21. Callitris roei [n=26, m=21, h=9, 1829–1997] Callitris roei growing in ‘mallee’ and ‘Melaleuca
thicket’ with some taller eucalypts in the background, near Harrysmith in SW Western Australia. Photo
credit Aljos Farjon.

Map AU-22. Callitris drummondii [n=29, m=27, h=10, 18??–

1997]

Map AU-23. Callitris endlicheri [n=73, m=68, h=13,

18??–1997]

Callitris endlicheri is a medium size tree up to 20 m

tall. It has an extensive range in eastern Australia,
occurring in and on either side of the Great Divid-
ing Range. Its habitat is more or less open eucalypt
forest and woodland, scrubland and grassland. The
trees are usually found on shallow, rocky soils, on
rock outcrops or in rock crevices where there may be
less competition from larger trees. This species also
invades overgrazed and then abandoned pastures and
is often found in disturbed terrain, such as road cuts
and road verges. In such places it may be accompa-
nied by alien, invasive plants like pines (Pinus) and
brooms (Cytisus scoparia). Although a native tree, it
is opportunistic like these invasives and takes advan-
tage of the additional dynamic conditions imposed on
the landscape by human activity. Its natural distribu-
tion is therefore likely to expand and its abundance to
increase in future.
358 australasia

Map AU-24. Callitris baileyi (black) [n=20, m=19, h=6, 1905–

2004]; C. muelleri (red) [n=26, m=22, h=10, 1800–1997]

Callitris baileyi is relatively rare, with main localities in

SE Queensland (Blackbutt Range, Bunya Mountains,
Coast Range, Leichhardt Highway, West Moreton),
and NE New South Wales (Clarence River, Killarney,
Tabulam). Its habitat is Eucalyptus forest, in which
it often occupies rocky outcrops of basalt, granite or
deposits of volcanic ash. It has therefore a scattered
distribution, occurring in small, often isolated sub-
populations. Callitris muelleri is a dwarfed tree or a
shrub usually not exceeding 4.5 m tall. It occurs from
near the coast up to the heights of the Blue Moun-
tains E of Sydney, at altitudes between 500m and 1050
m. Sandstone plateaux and escarpments or rocky hills
with scrub vegetation dominated by Myrtaceae and
Proteaceae and subjected to frequent wildfires are
its preferred habitat. Like other species in the genus,
it seems to benefit from soil disturbance and can be
seen congregating alongside dirt roads where the sur-
face has been scraped to the sides. The seed cones of
this species are large, serotinous and tenacious, protecting the seeds in case of fire. When the
vegetation is burned off, the cones on stems and branches open, releasing the seeds on to bare
ground fertilized with ashes.

Callitris muelleri on a dirt road verge in the Blue The serotinous seed cones of Callitris muelleri.
Mountains, NSW. Photo credit Aljos Farjon. Photo credit Aljos Farjon.
 australasia 359

Map AU-25. Callitris monticola [n=15, m=13, h=9,

1904–1997]

Callitris monticola is a dwarf tree (sometimes a shrub)

occurring in low eucalypt woodland, where it is virtu-
ally confined to outcrops of eroded boulders and rock
ledges. The rock types are a coarse granite (adamellite),
trachyte or less often sandstone. Here it can escape
both competition and forest fires as such places can
only support sparse and low vegetation. The altitudi-
nal range is from 560 m to 1360 m. This species has
a limited range in the low mountains on either side
of the border between Queensland and New South
Wales. The localities are separated and can be diffi-
cult to find as they often lie in inaccessible terrain.
In some previously known localities it has not been
found recently. Increased frequency or intensity of
fires are the main threat to this rare species, which is
now listed as Endangered (EN) (up from Vulnerable)
on the IUCN Red List.

Photograph of trees of C. macleayana in the Herberton

Map AU-26. Callitris macleayana [n=26,
m=22, h=9, 18??–1997]
Range near Cairns, northern Queensland. The natural
vegetation is a mosaic of wet sclerophyll forest on slopes
and ridges and rainforest in valleys, with C. macleayana
mostly occupying the higher ground. These trees are ca.
25 m tall and grow together with many species of angio-
sperm trees. Photo credit Aljos Farjon.
360 australasia

Callitris macleayana is exceptional in the genus for several reasons. Morphologically is is distinct
from other species in having two types of leaves: scale leaves in whorls of three as in other species,
and acicular leaves in whorls of four. Both types can produce seed cones and are therefore adult
leaves, consequently there are cones with six and cones with eight valvate scales on the same
tree. The thick stringy bark is also unlike that of other species. While all other Australian species
occupy arid or semi-arid habitats, only this species occurs in rainforest and wet sclerophyll for-
est bordering rainforest. It is the tallest species, attaining nearly 40 m, but such large old trees
are now rare. Its distribution is confined to the east coast and disjunct. In Queensland it occurs
near Cairns in (sub-)tropical rainforest and also near the coast close to Brisbane, with a gap of
nearly 1500 km between these populations. In New South Wales, where it is more widespread, it is
found between Grafton and Maitland, in coastal rainforest remnants as well as in wet sclerophyll
forest (dominated by Eucalyptus spp.) on red-brown clay (dolorite) or loam further inland. Both
morphology and phylogeny indicate a primitive evolutionary position of this species in relation
to the other Australian species, but with possible closer relationship to the two species in New
Caledonia (Farjon, 2005a; Piggin & Bruhl, 2010). This suggests that C. macleayana is an ancient
relict species from times in Australia’s geological past when the climate was cooler and wetter. It
now shelters in rainforest near the coast and on moisture catching E slopes of mountains to an
altitude of ca. 1000 m.

Two species of Callitris occur in Tasmania, but neither is confined to that island, which has oth-
erwise a quite unique conifer flora. Of the two, C. rhomboidea is the most common and wide-
spread species, occurring also in E Victoria and on Kangaroo Island in South Australia, along the
coast of New South Wales, and as far north as SE Queensland. The other species, C. oblonga, is
uncommon; the two disjunct populations in New South Wales have been treated by Australian
botanists as subspecies of C. oblonga.

Map AU-27. Callitris rhomboidea [n=94, m=82, h=12,

18??–2008]

Callitris rhomboidea is common in scrublands and

open woodlands, especially in areas with thin, ‘skel-
etal’ soils over bedrock or in rocky places and on
low mountain ridges. In the Grampians Mountains
of Victoria dense stands of small trees are found on
ridges and highest slopes, but elsewhere it is usually
more scattered, occurring in ‘mallee’ or open eucalypt
woodland. In eastern Tasmania it is common also in
land used for sheep grazing, where it can be accom-
panied by invasive alien (European) shrubs as well as
native species. It is apparently an opportunistic spe-
cies that can invade ground vacated by competitors
after disturbance. In Tasmania it is confined to the
drier E side of the island and to the islands of the Furneaux Group. Much of its range lies within
the zone of moist temperate climate with more or less equal rainfall through the year, but the
populations of E Victoria and Kangaroo Island are in the winter rainfall zone. This also indicates
that C. rhomboidea is an adaptable species with quite a broad ecological tolerance.
 australasia 361

Map AU-28. Callitris oblonga [n=43, m=39, h=12, 18??–2009]

In contrast, C. oblonga has a more narrowly circum-

scribed habitat. It is riparian, i.e. it follows streams and
is rarely found more than 100 m from a perennial water
course. In Tasmania, these are lowland streams running
through land mostly converted to pasture. Here, the spe-
cies is threatend by clearance of natural vegetation, pol-
lution, livestock-grazing and invasive species. The best
remaining stands are now put under a recovery conser-
vation programme by the Government of Tasmania. In
New South Wales the southernmost population occurs
in the drainage of the Corang River at around 600 m alti-
tude. This is about equidistant between the populations
of Tasmania and those in the north of NSW. The latter
are often very small, sometimes only a few score plants,
occurring along creeks in the New England Range at higher altitude, around 1100 m. Here trees
and shrubs are stunted or dwarfed and even the seed cones are smaller, a fact that has given rise
to the subspecies name parva. However it seems better to see this as an ecotype due to adverse
environment. This species is listed as Vulnerable (VU) on the IUCN Red List.

Podocarpaceae

In this section, the species in the family Podocarpaceae occurring on the Australian continent are
mapped, with the exception of Sundacarpus amarus, which occupies two small areas in Queen-
sland but is widespread in Malesia; it is treated there (see map MA-59 on p. 326). This leaves
seven species of Podocarpus, Prumnopitys ladei and Pherosphaera fitzgeraldii. One species of Pod-
ocarpus also occurs in Tasmania and is mapped here; conifer species endemic to Tasmania are
treated together separately. No species of Podocarpaceae is adapted to semi-arid conditions, so
in Australia they are confined to the margins of the continent in proximity to the coast or to high
altitudes when somewhat further inland. Most have limited ranges, but two, Podocarpus elatus
and P. spinulosus are widespread in Eastern Australia. Nearly half of these species are shrubs,
occurring in more or less restricted habitats; the four tree species occur in rainforest along the
E coast of the continent, three of these are confined to the tropics.

Map AU-29. Podocarpus drouynianus [n=18, m=15, h=7,

1844–2006]

Podocarpus drouynianus is the only species in the fam-

ily Podocarpaceae that occurs in Western Australia. This
species is ‘out on a limb’ in the far SW corner of West-
ern Australia. Its relationship with P. spinulosus in Sec-
tion Spinulosus according to De Laubenfels (1985) was
recently confirmed in a phylogenetic study based on
molecular data (Biffin et al. inTurner & Cernusak, eds.
2011). Both species are unusual in the genus in having
adaptations to survive in fire-prone vegetation, which
prevails in much of Australia (Ladd & Enright in Turner
& Cernusak, eds. 2011). Podocarpus drouynianus is found
from Bunbury to Mt. Barker and in the Darling Ranges.
Its habitat is tall Eucalyptus forest on sandy soils, where
362 australasia

it is a low shrub suckering from root stock, forming patches. This ability helps it to regenerate after
forest fires; it is also relatively immune to man-made disturbance such as logging. Yet it requires
the microclimate of the forest in the long term and does not occur outside the forest area.

Map AU-30. Podocarpus
spinu­losus [n=29, m28,
h=6, 1803–2004]
Podocarpus spinulosus under trees of Banksia aemula on North Strad-
broke Island, Queensland. The trunks of the trees are blackened by
fire, demonstrating that P. spinulosus can survive by rapidly resprout-
ing from root stock, taking advantage of the clearance of competing
vegetation. Photo credit Aljos Farjon.

Podocarpus spinulosus has a scattered distribution in Queensland and New South Wales, where it
occurs on sandy shores and islands as well as in locations further inland on soils derived from
sandstone deposits, such as in the Blackdown Tableland of Queensland. Its altitudinal range is
from 1 m to 900 m. It is a straggling or creeping shrub, commonly in the understorey of low wood-
land or mallee with Eucalyptus spp., Acacia spp., Casuarina spp. and Proteaceae as the dominant
tree species, or near the ocean in ‘heath scrubland’ with lower vegetation. These types of vegeta-
tion burn frequently and P. spinulosus can resprout after the above-ground parts are destroyed.
On the islands in Moreton Bay, Queensland, it grows on stabilized dunes covered in shrubs and
scattered eucalypts. Its layering habit allows it to spread over sometimes considerable areas.
Peculiarly, this species appears to be absent on Fraser Island, a large sandy island just N of the
mainland section of Great Sandy National Park E of Gympie, where it does occur. Along the coast,
that area seems to be the northernmost location of this species.

Map AU-31. Podocarpus elatus [n=38, m=37, h=5, 1802–2003]

Podocarpus elatus occurs in rainforest localities all along

the E coast of Australia from the Cape York Peninsula in N
Queensland to near Nowra in New South Wales; a disjunct
population is known from Arnhem Land in the Northern
Territory. It is fairly widespread and common in NSW and
SE Queensland, but further north it becomes rare and local-
ized, perhaps restricted to drier types of rainforest there. In
Flora of Australia 48 (K. D. Hill in McCarthy, ed. 1998) it is
“also reported from New Guinea” but this probably rests
on a misidentification, we have no herbarium records of
this species from there. Podocarpus elatus is a tree to 30 m
tall occurring in ‘complex mesophyll vine forest’, a type of
 australasia 363

rainforest with entire-leaved trees and abundant lianas and ­scattered palms, among which are
climbing rattan palms. There is ample rainfall but in the north there are dry spells from June to
September. The altitude range is from near sea level to ca. 1000 m and at higher altitudes it is
often confined to river banks.

Map AU-32. Podocarpus grayae [n=24, m=24,

h=6, 1821–2001]

Podocarpus grayae at Lake Eacham, Crater Lakes

National Park, Queensland. Photo credit Aljos Farjon.

Podocarpus grayae is distributed in N Queensland near the Pacific coast and, like P. elatus, also
has a disjunct population in Arnhem Land, Northern Territory. This is also a rainforest tree and
in N Queensland it is more common than P. elatus, often occurring on sand directly behind the
mangrove-covered tidal zone. It is most common inland along streams and occurs on mountain
ridges to an altitude of 750 m. Trees can attain 30 m as components of the complex notophyll
vine forest, where they are usually mixed in with many other tree species. The apparent gap in the
distribution on the Pacific coast of the Cape York Peninsula between Cooktown in the south and
the Stewart River in the north is also manifest in the map of P. elatus (map AU-31 on p. 362 and
no other species of Podocarpaceae have been found there. This is so because rainforest is almost
absent along this stretch of the coast, limited to a few tiny patches only. Instead, in areas with
natural vegetation an open landscape with Eucalyptus and termite hills is the dominant feature
here, due to frequent fires and seasonal flooding.
364 australasia
Map AU-33. Podocarpus smithii Map AU-34. Podocarpus disper- Map AU-35. Prumnopitys ladei
[n=17, m=17, h=4, 1936–2007] mus [n=41, m=40, h=5, 1915–1999] [n=21, m=21, h=5, 1908–2001]
Prumnopitys ladei in rainforest on Mt. Lewis NW of
Cairns, Queensland. This rare species is one of nine
species in a genus that has a very disjunct distribution,
with species in Australia, New Zealand, New Caledonia,
Central America and South America. This distribution
and its biogeographical implications are discussed in
more detail in the chapter South America. Photo credit
Aljos Farjon.
These three species of Podocarpaceae
have limited ranges in the same area of
N Queensland near Cairns. Podocarpus
smithii and P. dispermus are both present
in the Atherton Tableland; P. smithii and
Prumnopitys ladei are both on Mt. Lewis.
Podocarpus dispermus is the only species
in the Bellenden Ker Range to the SE of
the Atherton Tableland and Prumnopitys
ladei is also found on Mt. Spurgeon and
a few other localities nearby. Podocarpus
dispermus occurs at lower altitudes than
the other two (below 750 m); P. smithi
and Prumnopitys ladei are found between
900 m and 1400 m in montane rainforest.
Both Mt. Lewis and Mt. Spurgeon are high
granitic outcrops rising above the Ather-
ton Tableland, which consists of sand-
stone and volcanic formations. Especially
with the latter two species, populations
are small and these trees can be consid-
ered rare. While natural forests in the
region are now mostly protected, this is
a recent change in policy and logging still
happened in some areas in the 1980s. It
is assumed that the abundancy of mature
trees has as a result declined, but that the
geographical distribution shown by the
maps has largely remained unaltered.
Prumnopitys ladei is listed as Vulnerable
(VU) on the IUCN Red List.
 australasia 365

Map AU-36. Pherosphaera fitzgeraldii [n=10, m=10, h=4,

1881–1998]

This is one of two species in this genus, formerly but

incorrectly known as Microstrobos. The other spe-
cies is endemic in Tasmania and are treated with the
other endemic conifers of that island. Pherosphaera
fitzgeraldii is extremely localized, as it is found only
near some of the waterfalls that cascade down from
the Blue Mountains in New South Wales. The popula-
tion at Wentworth Falls is best known, another local-
ity nearby is at the Katoomba Falls, both the upper
and lower falls have tiny populations. Pherosphaera
fitzgeraldii is a decumbent shrub that only occurs in
the spray zone at the lower part of waterfalls with permanent clean water supply.
At the beginning of the twentieth century this species was reported from “most of the chief
falls on the Blue Mountains”, but it has suffered a sharp decline and is now down to fewer than
500 mature plants at the waterfalls mentioned. Although mostly situated within a national park,
water pollution caused by urban developments upstream, habitat degradation and invasive spe-
cies have all played a part in this decline. It is listed as Critically Endangered (CR) on the IUCN
Red List.

Map AU-37. Podocarpus lawrencei [n=76, Podocarpus lawrencei among rocks on Cradle Mountain, Tasma-
m=76, h=11, 1804–2008] nia. Photo credit Aljos Farjon.

Podocarpus lawrencei is an alpine or subalpine decumbent shrub occurring both in mailand Aus-
tralia and in Tasmania. In New South Wales, A.C.T. and Victoria it is present in the ‘Australian
Alps’, the highest and most southern part of the Great Dividing Range, where it is found to 2030 m.
In Tasmania it also occurs in the mountains, but these are lower; due to a much cooler climate
this species can there grow down to 450 m, but it is mostly found from 1000 m to 1350 m a.s.l.
These shrubby podocarps usually occur above the tree line in heath-like vegetation or herbaceous
grassland and among rocks, where snow covers the ground for variable periods of time in winter.
On the mainland this species can be part of the understorey in low sclerophyll forest dominated
by Acacia, Eucalyptus and Telopea and will then occasionally attain a height of 4 m. It is locally
common, or even abundant, but remains a lesser component of the vegetation.
366 australasia

Tasmania

Geography

Tasmania is a large island situated to the SE of the Australian continent, separated from the state
of Victoria by Bass Strait at a distance of 250–300 km. It has an area of 67,800 km² including its
satellite islands and is mostly mountainous, but with modest elevation (highest point Mt. Ossa at
1617 m) and with lowlands in the eastern part of the island. Situated approximately between 40°
30’ S and 43° 30’ S in the cold Southern Ocean its climate is markedly cooler than in the southern
parts of the continent and on the west coast a perpetually cool and wet climate prevails, with
annual precipitation exceeding 3000 mm. Snowfall is possible in the mountains at any time of
the year (although there is no permanent snow cover) and summers are cool and cloudy much
of the time. On the E coast the climate is much drier and more or less similar to parts of coastal
Victoria, with more seasonal rainfall. Tasmania was the last part of Australia to separate from
Antarctica at the end of the Eocene, but it remained connected with Australia during the Pleis-
tocene glacial maxima, when sea level dropped more than 100 m. It has a high level of endemism
notwithstanding its proximity to the continent, between 12–15% of plant species, and this is even
more pronounced in conifers. Four genera and eight species are endemic out of a total of 13 spe-
cies, and Phyllocladus is present with one species here, but absent from mainland Australia. The
fossil record shows an extraordinary diversity of conifers present during the Late Cretaceous and
Tertiary, with 24 genera so far identified (Hill & Brodribb, 1999) many of which occur at present
outside Australia (Farjon, 2008, p. 173). Tasmania apparently was a ‘hub’ for conifer distribution
from the Late Cretaceous until it became separated by a widening ocean from Antarctica. Espe-
cially the fossil taxa discovered here establish links with New Zealand, New Caledonia and Fiji,
Australia, New Guinea and South America. Its conifer diversity, despite the extinctions, is still
extraordinary in relation to the landmass of the island and in comparison to the nearby Austra-
lian continent.

Athrotaxis

The genus Athrotaxis (Cupressaceae) is endemic to Tasmania and is here considered to have three
species, A. cupressoides, A. selaginoides and A. × laxifolia. The latter is considered to be of hybrid
origin with the two other species as parents and Australian botanists tend to treat it as a recur-
rent F1 hybrid, but the evidence for this is not conclusive and it appears to breed true to type in
cultivation. There is evidence in the fossil record of Athrotaxis in South America and New Zealand
(Farjon, 2008, p. 173) and therefore it is likely that the three taxa today limited to Tasmania are
relicts of a more diverse and widespread genus of Gondwanan origin. This genus (formerly clas-
sified in the family Taxodiaceae which has been subsumed in Cupressaceae) is phylogenetically
a representative of a basal ‘grade’ of mainly relict taxa and the only representative of this loosely
defined group of genera in the Southern Hemisphere today. Ecologically if not phylogenetically
it has equivalents in the family occurring both in South America (Austrocedrus, Fitzroya, Pilgero-
dendron) and in New Zealand (Libocedrus) and two of these have also been found as fossils in
Tasmania. All are members of Cupressaceae mostly adapted to a cool and wet mountain climate
influenced by the southern oceans, today with only 1–5 species left per genus.
 australasia 367

Map AU-38. Athrotaxis cupressoides [n=41,

m=40, h=12, 18??–2009]

This is the most abundant species in the

highest parts of the island, occurring at
altitudes between 900 m and 1300 m. Sub-
stantial populations occur on the Central
Plateau, the Great Western Tier and on
mountains W of these e.g. around Cradle
Mountain and Mt. Field. In these areas it
can form dense stands, often with abrupt
transitions to ‘Buttongrass moorland’ or
dwarf scrub, or congregate around lake
and ‘tarn’ shores. At its lower altitude
range it merges into forest with Eucalyp-
tus and Nothofagus, but becomes more
scattered and is often replaced by A. selaginoides. Athrotaxis cupressoides often grows on peaty
soil and in dry spells this in conjunction with the tussock grassland can expose it to fires that
kill the trees. Large fires occurred in 1960/61 on the Central Plateau, destroying large parts of the
population. Grazing by introduced animals, especially rabbits, prevents regeneration. This species
is classified as Vulnerable (VU) on the IUCN Red List.

Map AU-39. Athrotaxis selaginoides [n=42, Athrotaxis selaginoides on the shore of Dove Lake, Tasma-
m=42, h=13, 18??–2008] nia. Photo credit Aljos Farjon.

Athrotaxis selaginoides has a wider distribution in W Tasmania than A. cupressoides, but the map
based on herbarium collections presented here has no records from the SW part of the island.
It occurs there according to Brown & Hill (in Farjon & Page, 1999, map on p. 68) but no records
have been found in the institutional herbaria consulted (BM, CANB, CBG, CGE, E, HO, K, NY, S).
It is presumably rare in that area. This species is more or less common in the forest zone above
Eucalyptus tall forest but below the subalpine zone, where this species if it occurs will retreat
to sheltered localities and on lake shores or along streams. Its altitude ranges from ca. 400 m
to 1200 m. As a forest tree it can reach large size and has consequently been subject to logging,
which may account for its scarcity in some areas. Most remaining trees are now in protected
areas and it is no longer logged. Sensitivity to forest fires has meant further decline, as these fires
have increased beyond natural frequencies and intensity, especially in forests affected by logging
operations. This species is classified as Vulnerable (VU) on the IUCN Red List.
368 australasia

Map AU-40. Athrotaxis ×laxifolia. [n=13,

m=13, h=6, 18??–1998]

This putative nothospecies is the least

common of the three taxa in the genus
Athrotaxis, with a scattered occurrence
most commonly in the northern part of
the range of A. cupressoides. It is usu-
ally but not exclusively found where
the two putative parent species occur,
giving some support for the hybrid
hypothesis. Genetic research which
used random amplified DNA markers
also supported this origin (Isoda et al.,
2000a) but was inconclusive in terms
of the taxonomic status of the hybrid.
This taxon is listed as Vulnerable (VU)
on the IUCN Red List.

Other Conifers

Map AU-41. Phyllocladus aspleniifolius

(Phyllocladaceae) [n=90, m=90, h=12,
1792–2008]

Phyllocladus aspleniifolius is one of

four species in the genus Phyllocla-
dus and the only one that occurs in
Australia. The interesting distribution
of the genus is discussed under New
Zealand (Map AU-64 on p. 384 where
two of the four species occur. The Tas-
manian species is widely distributed
on the island, predominantly in the
mountainous western half but also
with locations on some of the islands
on the E coast and on high points in
the NE. There is again a paucity of
herbarium collections from the SW of the island, although it is common there as well (Brown
& Hill in Farjon & Page, 1999, map on p. 70). On the wet W coast it descends down to sea level
and in the interior it ascends to the alpine zone at around 1200 m. It is a tall forest tree among
eucalypts at middle altitudes; from about 750 m upwards it is a constituent of mixed forest where
besides Eucalyptus coccifera two species of Nothofagus are co-dominant, as well as the conifers
Athrotaxis and Phyllocladus. Still higher it becomes a stunted tree in open woodland and heath-
like vegetation dominated by shrubs and perennial herbs.
 australasia 369

Map AU-42. Lagarostrobos franklinii [n=52,

m=49, h=15, 1845–2004]

Lagarostrobos franklinii is a member of

Podocarpaceae and here considered to
be the only species in the genus. Its
nearest relative is Manoao colensoi in
New Zealand; by some botanists the
generic split is not accepted, under
which concept there would be two spe-
cies in Lagarostrobos. Lagarostrobos
franklinii is a riparian tree, i.e. it occurs
almost always on the banks of rivers
and shores of (lowland) lakes. Excep-
tions are found on the extremely wet W
coast, most famously on Mt. Read, where
an ancient tree long ago started layering, spreading its stems across a large area. On this mountain
it occurs at ca. 1100 m, but along the rivers it is a lowland tree, from just above sea level to ca. 160
m. It forms narrow groves lining the rivers, but can be mixed with Nothofagus cunninghamii,
Eucryphia lucida, Anopterus glandulosus and tree ferns. Away from the W coast in upland terrain
these riparian forest strips are flanked by forest dominated by Eucalyptus obliqua. Huon Pine was
once the most important timber tree of Tasmania, and logging along the rivers has removed
almost all the large trees. Small and often curved trees were left, and these together with re-
sprouting and other re-growth have saved the species, which is now assiduously protected. A few
isolated groves of large trees escaped attention from the loggers; usually these were some distance
away from larger rivers or streams on which the logs could be transported.

Map AU-43. Pherosphaera hookeriana (Pod-

ocarpaceae) [n=36, m=36, h=12, 1847–2011]

Pherosphaera hookeriana is one of two

species in this genus; P. fitzgeraldii
occurs in New South Wales (map. AU-36
on p 365). It was formerly known as
Microstrobos niphophilus but this name
is incorrect under the rules of botanical
nomenclature. It forms an erect shrub
to 2.5 m tall. This species has a limited
distribution in the higher parts of Tas-
mania. It is locally common in subal-
pine vegetation above 1000 m but has
been found as low as 700 m a.s.l. It is
frequent in wet moors and often borders
the lakes and ‘tarns’ that are numerous
in the mountains. Most of the herbarium collections used to map this species have been made in
Mt. Field National Park and vicinity, but it is common also in other areas, e.g. Cradle Mountain-
Lake St. Claire National Park.
370 australasia

Microcachrys tetragona with ripe cones. Photo

credit Aljos Farjon

Map AU-44. Microcachrys tetragona (Podocarpaceae)

[n=57, m=57, h=12, 1893–2008]

Microcachrys tetragona is one of the smallest conifer species, creeping over ground and on rocks
in the mountains of western Tasmania. The species is a monotypic dwarf conifer widely distrib-
uted on the western mountains of Tasmania. Herbarium collecting has apparently focused on
two national parks, Mt. Field and Cradle Mountain-Lake St. Clair, and some other high points
in the central and western mountains, as well as in the south, are represented. This small plant
creeps over rocks and peaty ground and is only 10–30 cm tall. It is restricted to alpine vegetation
at altitudes above 1000 m and can be abundant in many localities with suitable habitat.

Map AU-45. Diselma archeri (Cupressaceae) [n=42,

m=40, h=11, 1841–2008]

Diselma archeri is a monotypic shrub or rarely

a small tree to 4 m tall occurring in the moun-
tains of W Tasmania, where it is widespread
and common in several locations. It is a sub-
alpine to alpine species predominantly found
above 1000 m but its lowest station known
from a herbarium collection is at 570 m on
the Nive River. It grows often together with
Pherosphaera hookeriana and although these
two coniferous shrubs look very similar, they
belong to different families and can be identi-
fied by their female cones, which are tiny and
inconspicuous and, in both species, only present on female plants. Another, but more difficult
character to accurately observe due to minute leaf size is phyllotaxis, decussate in Diselma and
spiral in Pherosphaera.
Misidentification is therefore likely to occur and distribution data should preferably be based
on verifiable herbarium specimens, as in this Atlas.
 australasia 371

Diselma archeri near Cradle Mountain in typical habitat: high elevation rocky terrain dotted with ponds
and small lakes (‘tarns’) and with a mixture of ‘heath’ vegetation and shrubs and low trees, many of which
are conifers. Photo credit Aljos Farjon.
372 australasia

New Zealand

Geography

New Zealand is an isolated group of two large and several small islands situated at temperate
latitudes (between ca. 34° S and 47° S) in the SW Pacific Ocean at approximately 2000 km from
the nearest continent Australia. The total land area is ca. 267,000 km². North Island extends with
a narrow peninsula (Northland) into the subtropics, while South Island and its southern satel-
lite Stewart Island reach into the cold temperate climate zone. High mountains are present on
both main islands. In North Island these are volcanoes and form more or less isolated massifs or
mountains culminating in Mt. Ruapehu (2797 m) in the centre of the island. In South Island the
Southern Alps extend along much of the W coast and are in part glaciated, the highest peak here
is Aoraki (Mt. Cook) at 3754 m. There are also extensive lowlands, notably on the E side of South
Island, and large bays dotted with small islands. The climate is moist to wet maritime, but E of
the Southern Alps there is a marked rain shadow effect causing steep gradients in precipitation.
The islands are uplifted parts of a subcontinent that was joined with Antarctica and Australia but
became submerged. Widening of the Tasman Basin increased the separation, while northwards
connections persisted or were created, e.g. the Lord Howe Rise and Norfolk Ridge, to now sub-
merged or nearly submerged isolated islands such as Lord Howe, Norfolk Island and perhaps New
Caledonia. New Zealand has been separated from other landmasses since the Late Cretaceous
about 80 Ma and is now one of the most isolated major landmasses on Earth.

Conifers in New Zealand

Families Genera Species Species + infra Countries with Area of occupancy Endemic taxa
conifers in km²
4/8 10/70 19/615 20/794 1/1 19,000/267,000 20
Numbers after ‘/’ are global or country totals.

The remoteness of New Zealand for such a long time has caused a unique fauna and flora to
evolve. Before humans arrived only about 1000 years ago there were no mammals except a few
bats and seals and many niches elsewhere filled by mammals were occupied by birds, many of
these became flightless. Evolution in isolation led to high levels of endemism, ca. 80% of vascular
plants, 90% of freshwater fish and all amphibians and reptiles. All taxa (19 species + 1 variety) of
conifers are endemic, too. At the generic level, only two genera out of 10 are endemic: Halocarpus
and Manoao, and there are no endemic conifer families. At these higher taxonomic levels, links
are with other species mostly distributed in various Gondwanan lands, e.g. New Caledonia, Fiji,
Tasmania and South America, suggesting vicariance. Immigration mainly from Australia has also
contributed to the flora of New Zealand, and some species of Podocarpus now endemic to New
Zealand may have an Australian ancestor.
The late arrival of humans in New Zealand meant that their impact on ecosystems was pro-
found and rapid. Extinction especially of flightless birds followed quickly and the forest vegeta-
tion of the lowlands was altered in probably 50% of the area before Europeans arrived. In the two
most recent centuries the latter have altered the landscape below the higher mountains perhaps
more drastically than in other regions that they colonized. The forests were severely reduced in
area and fragmented to isolated patches, especially in the lowlands. Agricultural and urban land
uses have made these changes irreversible, at least for the duration of European civilization. More
contiguous native forest now exists mainly in the high mountains, especially in South Island. By
the time of the National Forest Survey in 1955 ca. 62,000 km² of native forest was left, the greatest
part of it in mountain areas but with numerous small remnants elsewhere. Especially in the low-
 australasia 373

lands and along forest fringes much of this is secondary forest or scrubland now slowly reverting
back to mature forest. About 90% of native forest is now in protected areas or its native trees are
under protective legislation if on private land. It is quite difficult to assess what this large scale
alteration of the landscape in a relatively short time has meant for the present distribution of
conifers. Certainly where forest has been turned to pasture for sheep or cattle they no longer
occur, although even there solitary trees may have been left standing and, if recorded in a her-
barium specimen, these will give a dot on the map. The scattered distribution of large and small
forest reserves has a similar effect. Absence or presence on the distribution maps of New Zealand
conifers are therefore not sufficient indicators of present forest cover and the distribution pat-
terns shown must be interpreted with caution in that context.

Map AU-46. The distribution of conifer spe-

cies in New Zealand.

Conifers in New Zealand occur in several

forest types. In the far N of North Island
Kauri forest is dominated by Agathis
australis (Araucariaceae) and has a mix-
ture of several species of Podocarpaceae
and angiosperm trees. This is lowland
subtropical to warm temperate forest. A
forest type mostly reduced to remnants
occurs in the lowlands elsewhere and is
known as Podocarp-mixed broadleaf for-
est. Often the dominant tree species is
the conifer Dacrydium cupressinum
(Podocarpaceae) and several other spe-
cies of that family occur here, too, besides
various angiosperms. At high altitude a
mixed forest type dominated by Liboce-
drus bidwillii (Cupressaceae) also has various other conifers frequently present. Dacrycarpus
dacrydioides is often dominant in swamp forests and other areas with high water tables and due
to unsuitability for agriculture these have often been spared the axe. Finally, conifers occur and
are often prominent at high altitude in subalpine scrubland and some species are limited to this
habitat. The conifer flora of New Zealand is relatively modest in numbers of taxa, but notable for
its endemism and important ecologically.

Conifer forest in Northland, North

Island, New Zealand. Species present
in this photograph are Dacrycarpus
dacrydioides (left), Podocarpus totara
and Phyllocladus trichomanoides (right)
and Agathis australis (tall tree in back-
ground). The only non conifer is the
(unidentified) tall shrub in the lower left
corner. Those of us only familiar with
conifer forests in the Northern Hemi-
sphere may have difficulty recognizing
conifers in this picture; their habits as
well as the forest structure are totally
different. Perhaps this is a glimpse of
what the forests of Gondwana were like
80 Ma. Photo credit Aljos Farjon.
374 australasia

Agathis

Map AU-47. Agathis australis [n=61, m=61, h=7, 1826–2010]

The famous Kauri pine of New Zealand, Agathis aus-

tralis (Araucariaceae), has a limited distribution in
North Island. It occurs from Kawhia Harbour and
Maketu in the south to North Cape and is present on
many of the islands as well as on the mainland. Despite
the massive logging that occurred a little more than a
century ago the extent of occurrence (EOO), indicated
on the map by the peripheral herbarium collection
locations, is nearly equal to its historical range. The
decline has been in the area of occupancy (AOO), esti-
mated to have gone from 1,215,000 ha before logging
to fewer than 14,000 ha at present. The scattered dis-
tribution of numerous forest remnants large and
small, now almost all with a protected status, accounts
for the current EOO. An estimated 80,000 ha of these forests are suitable habitat for the Kauri
pine and under proper management this potential increase of the AOO of Agathis australis can
be realised in future. Perhaps ironically, large scale disturbance of the forest is a prerequisite for
Kauri pine regeneration (Ogden & Stewart in Enright & Hill, eds. 1995), as without these events
other trees (angiosperms in particular) will out-compete Agathis australis and eventually replace
it. Old trees survive for many centuries as emergents rising above a tree canopy under which its
seedlings have little chance of becoming trees themselves. Cyclones and fires would be the natu-
ral agents of canopy destruction and open large patches of forest, while the massive boles of the
largest Kauri pines resist the destruction, making them the seed trees of the next generation. It is
likely that many of the present forest reserves are too small for these processes to unfold and
unless they can be connected or otherwise enlarged, some form of active management will be
required to ensure ‘natural’ regeneration. Kauri pines are not evenly distributed in the remaining
forest reserves for an additional ecological reason common with conifers known as niche parti-
tioning. Better able to cope with nutrient deficient soils than most angiosperms, they tend to be
more numerous on ridge crests in hilly terrain, from where leaching and downward erosion have
transported these essential minerals downslope. In small reserves with this kind of topography
the Kauri pines may persist longer without interference.

Agathis australis in Trounson Kauri Park,

North Island, New Zealand. These two boles
are ca. 1.5 m diam. and straight and tall. In
an initial phase, trees grow fast in length
but not in girth (‘rickers’) to keep abreast of
the canopy or if possible overtop the com-
petition of tree ferns (seen here) and angio-
sperms. These two trees are in a second
phase, developing a spreading crown and
increasing girth. Large trees may exceed 5 m
diam. with the thickest extant tree known
as Te Matua Ngahere in Maori (Father of
the Forest) measuring 5.2 m, but the larg-
est recorded specimen, known as the Great
Ghost, measured 8.54 m. It was destroyed by
fire in 1890. Photo credit Aljos Farjon.
 australasia 375

Libocedrus

The genus Libocedrus (Cupressaceae) is distributed in New Zealand (2 species) and in New Cale-
donia (3 species). Phylogenetic analysis has shown Pilgerodendron uviferum, a species from S Chile
and adjacent Argentina, to belong to the clade containing four of the five species of Libocedrus
(Leslie et al., 2012) a result earlier obtained by Gadek et al. (2000). In cladistic classification this
result would sink Pilgerodendron into Libocedrus, but the South American species is morphologi-
cally distinct and geographically highly disjunct. An inferred divergence time for Pilgerodendron
somewhere in the Late Oligocene to Miocene resulting from the most recent DNA analysis (Leslie
et al., 2012) invokes a dispersal event across the South Pacific as by that time land connections
between Australasia and South America via Antarctica did no longer exist. There are as yet few fos-
sils that can help elucidate this question. Libocedrus is known from Palaeocene to Miocene strata
in Australia (New South Wales and Tasmania) and Austrocedrus (extant in South America and
sister to the Libocedrus clade) is known from the Oligocene to Miocene in Tasmania (Hill & Bro-
dribb, 1999). Some link across Antarctica involving this clade seems likely, but does not solve the
late divergence time unless we assume that Libocedrus existed in Antarctica (and perhaps South
America) until the start of the Miocene, when South America finally separated from Antarctica.

Pilgerodendron Libocedrus

Map AU-48. The distribution of two closely related genera in Cupressaceae: Pilgerodendron in South Amer-
ica and Libocedrus in New Zealand and New Caledonia. Australia and Tasmania, where none of these
conifer genera occur now, has yielded Libocedrus in the fossil record of the Paleogene (early Cenozoic),
suggesting a link via Antarctica.

Map AU-49. Libocedrus bidwillii [n=82, m=81, h=7, 18??–

2002]

Libocedrus bidwillii is distributed both in North Island

and in South Island. It occurs in mountains from 250
to 1370 m altitude and is a co-dominant or dominant
tree in montane to subalpine evergreen rainforests of
a mixed type with conifers and angiosperms. Its lon-
gevity, 800–1000 years, exceeds that of any angio-
sperms present, indicating dependency for regeneration
on stand-replacing episodic disturbances such as
cyclones or fires. Several other conifers that grow with
it, such as Halocarpus biformis, Phyllocladus tricho-
manoides var. alpinus, and Podocarpus cunninghamii
have shorter life spans. At lower altitude, Dacrydium
376 australasia

cupressinum tends to dominate and Libocedrus is a minor component of the forest. Frequent
angiosperm trees at higher altitude are Metrosideros umbellata, Nothofagus solandri, Quintinia
acutifolia and Weinmannia racemosa, and Cordyline indivisa is a prominent shrub in this forest
type. Often the forests dominated by Libocedrus bidwillii (‘cedar forest’) grow on peaty, water-
saturated soil or in drainage systems surrounded by moorland. The cluster of localities in the
centre of North Island and those in Westland of South Island mostly represent these wet and
acidic habitats.

Libocedrus bidwillii in the Kaimanawa Mts. of

North Island, New Zealand. At this location it
forms nearly pure ‘cedar forest’ with a dense
undergrowth of non conifer shrubs. Nearly all
trees are old or senescent and very little regen-
eration takes place due to the dense shrub layer;
the ‘wait’ is for a forest fire to sweep this out of
the way. Photo credit Aljos Farjon.

Map AU-50. Libocedrus plumosa [n=66, m=63, h=8,

18??–2006]

The habitat of Libocedrus plumosa differs from that of

the previous species, although it also occurs in mixed
conifer-angiosperm forest. It is confined to the low-
lands but can occur to 600 m atlitude. Its distribution
coincides with the range of Agathis australis (see map
AU-47 on p. 374) but in most Kauri forests it is not
common. It is rather a constituent of a more diverse
lowland forest type, known as Podocarp-mixed broa-
dleaf forest, in which conifers form a mixed canopy
with angiosperms. Undisturbed forest of this type can
have as many as eight conifer genera (and species) on
a single hectare (Ogden & al., 1993). A few outlying
locations of L. plumosa are known, most notably in the
northernmost part of South Island around Whanganui
Inlet, where it occurs in mixed forest at low altitude
from near sea level to around 100 m, and S of there on
‘Cedar Ridge’ associated with Nothofagus.

Podocarpus

The large genus Podocarpus (Podocarpaceae) is represented in New Zealand with four species.
Two of these are shrubs (or rarely a small tree in one case), the other two are trees, of which only
P. totara can reach large size. Three species are spread more or less widely over the three main
islands (including Stewart Island in the far south) and one, P. acutifolius, is limited to South Island.
Three of the four species were included in a phylogenetic analysis based on molecular (DNA
sequence) data and are part of a clade with P. gnidioides and P. lawrencei (Biffin et al. in Turner
& Cernusak, eds. 2011). Podocarpus gnidioides occurs in New Caledonia (map AU-109 on p. 415 and
 australasia 377

P. lawrencei in Australia (map AU-37 on p. 365 It is likely that P. acutifolius belongs here, too as
its morphology is in several characters similar to P. totara. Whereas the fossil record has not
yielded similar links as described for Libocedrus (p. 375 we can infer a similar biogeographical
history, with links to at least Australia (+ Tasmania) and New Caledonia. The estimated diver-
gence time for this clade is sometime in the Neogene, i.e. later than 23 Ma. As both New Zealand
and New Caledonia were by then a long way from Australia, dispersal events must be responsible
for this distribution of related species. The most likely direction of migration would have been
from a drying Australian continent, with podocarps mostly pushed to the E coast fringe, and by
means of seeds carried by birds across the widening ocean to the islands.

Map AU-51. Podocarpus cunninghamii [n=115, m=114, h=6,

1841–2010]

Podocarpus cunninghamii (syn. P. hallii) is widespread

all over New Zealand and occurs from near sea level
to around 1000 m in mixed evergreen forest or forest
dominated by conifers. It is a moderately tall tree to 20
m but despite this it can be dominant in certain loca-
tions where the forest canopy remains low. This spe-
cies is easily dispersed by birds and can establish itself
in open areas of secondary forest, scrubland or even
grassland. Below 600 m it can co-exist with P. totara
and is thought to occasionally hybridize with it, above
this altitude it is the only tree forming species of the
genus. In South Island dense stands of Nothofagus
solandri are its most serious competitor, and it can only prevail as an emergent in lower forests
at high altitude or exposed ridges.

Map AU-52. Podocarpus totara, [n=108, m=105, h=6, 1829–2010] The Pouakani totara near Mangapehi, North
Island is the largest specimen tree of this species today. It was left by loggers as a curiosity. A photograph
by J. T. Salmon in 1979 (Salmon, 1980; 1986) shows this tree standing in a more or less open landscape, but
when this photograph was taken in 2002 it stood in a young secondary forest. Photo credit Aljos Farjon.

As with P. cunninghamii, P. totara occurs all over New Zealand, but it is more common in North
Island than in South Island and Stewart Island. It is a constituent or sometimes co-dominant tree
of lowland evergreen Podocarp-mixed broadleaf forest. Mature, undisturbed forests of this type
are species rich, with many tree ferns, angiosperm trees and abundant epiphytes, and several
378 australasia

s­ pecies of podocarpaceous conifers, sometimes with Libocedrus plumosa and in the far north with
Agathis australis added. Podocarpus totara can become a very large tree to 40 m tall and 3.6 m
diam. but such large trees are now very rare due to logging. Its longevity has been estimated at
around 900 years, indicating K-strategy ecology, with long cycles of regeneration to death. Defor-
estation of the lowlands may have had an impact on the distribution of this slow growing species
and it may have been more widespread in the past than the map now indicates.

Map AU-53. Podocarpus nivalis [n=92, m=91,

h=5, 1839–2006]

Podocarpus nivalis is an alpine or sub-

alpine shrub distributed in all the high
localities of New Zealand between 800
and 2500 m. On Stewart Island it occurs
down to 650 m a.s.l. It is common in open
scrubland and tussock grassland and can
spread vegetatively to form wide bushes
or patches. In rocky terrain it becomes
decumbent and although evergreen, snow
can cover it for several months in winter
at the highest altitudes. One would expect
an altitudinal gradient to occur, with the
southernmost populations at lower alti-
tudes than those on North Island, but the
available data from the herbarium speci-
mens do not confirm this except on Stew-
art Island, which has a very cool maritime
climate. This species will have been less
affected (but not unaffected: introduced
chamois and deer have caused erosion of
its habitat by overgrazing) by the human
footprint imposed on these islands and
assuming a near comprehensive cover-
age of herbarium collecting in the Conifer
Database the map may show the natural
distribution fairly accurately. Effects of
the warming of the climate during the
past 150 years or so are not yet apparent
(or have not been studied), but may have
a notable impact in the future.
Podocarpus nivalis creeping over rocks in Tongariro
National Park, North Island, New Zealand. Photo credit
Aljos Farjon.
 australasia 379

Map AU-54. Podocarpus acutifolius [n=36, m=36, h=4,

1875–2001]

Podocarpus acutifolius occurs in the lowlands and foot-

hills of the mountains in the northern part of South
Island, from Nelson in the north to Jacobs River in
the south. Its distribution appears to be limited to the
coastal area and lower slopes exposed to the moisture
coming in from the Tasman Sea. The altitude ranges
from near sea level to 650 m. Most often it is a shrub,
decumbent or erect, but occasionally a small tree to
9 m tall and a trunk to 40 cm d.b.h. Those tree forms
are allegedly hybrids between P. acutifolius and P. cun-
ninghamii or P. totara, both of which occur in the same area and beyond. Such hybrid identity, if
indicated with collected specimens, has been excluded from the database and this map, regard-
less if it is correct or not.

Prumnopitys

The genus Prumnopitys (Podocarpaceae) has nine species with an interesting, disjunct distribu-
tion of Gondwanan origin, currently with the highest diversity and greatest geographical range in
Central to South America. The geography of the genus is discussed in the chapter South America
(see map SA-5 on p. 432). Two species occur in New Zealand, discussed here. They have very simi-
lar ranges, spread across all three main islands, but more sparsely or scattered in South Island,
where they are mostly absent in the interior, unlike the situation in North Island. This may be
caused by the rain shadow effect imposed by the high mountain chain of the Southern Alps. Both
species are common constituents of the Podocarp-mixed broadleaf forest of the lowlands up to
800–1000 m, a temperate rainforest type rich in epiphytes and thriving only in areas with high
rainfall throughout the year.

Map AU-55. Prumnopitys ferruginea [n=116, m=113, Map AU-56. Prumnopitys taxifolia [n=124, m=120,
h=5, 1826–2010] h=8, 1769–2010]
380 australasia

On Stewart Island Prumnopitys ferruginea is found in monospecific stands with Pseudopanax

crassifolius in the understorey. Prumnopitys taxifolia may reach an age in excess of 1000 years
and is therefore a typical K-strategy tree: slow growing, long living, irregular regeneration con-
nected with forest disturbances that occur sporadically. Such trees can only survive naturally in
large, unmanaged forests where these natural processes can play themselves out, not in managed
patches of forest that are artificially kept in a pioneer phase dominated by young trees. The dis-
tribution pattern of these species at present may not, despite the protected status of native forest
remnants today, reflect their long term future.

Dacrydium

Map AU-57. Dacrydium cupressinum [n=81, m=78, h=5,

1770–2010]

Dacrydium cupressinum (Podocarpaceae) known as rimu

(Maori) in New Zealand, is a common and usually domi-
nant large conifer in Podocarp-mixed broadleaf forest of
the lowlands up to 700 m throughout New Zealand. This
species is most abundant in North Island and the north-
ern part of South Island, further south it is restricted to
locations on the coast and on Stewart Island. In Northland
of North Island it occurs in the Kauri forest with Agathis
australis as well as in its more typical forest type. These
forests are (warm) temperate evergreen rain forests with
year-round high precipitation and are multi-layered with
emergent conifers, a canopy of conifers and/or angiosperms, an understorey of shrubs, tree ferns
and palms, and a ground cover of ferns and mosses. Dacrydium cupressinum is a typical example
of a long-lived ‘catastrophic regeneration’ conifer (Ogden & Stewart in Enright & Hill, eds. 1995)
and the oldest trees known are from 900–1160 years. This species, when it becomes an emergent
forest giant, is particularly suitable for epiphytic growth and large trees can become complex
micro-habitats for a great number of species of plants, fungi and animals.

Dacrydium cupressinum in Tongariro National Dacrydium cupressinum in Tongariro N.P. heav-

Park, North Island, New Zealand. It is an emergent ily loaded with epiphytic plants, such as Astelia
tree in the forest. Photo credit Aljos farjon. solandri (Liliaceae), orchids and ferns. Photo credit
Aljos Farjon.
 australasia 381

Dacrycapus

Map AU-58. Dacrycarpus dacrydioides [n=106, m=102, h=5,

1826–2010]

Dacrycarpus dacrydioides (Podocarpaceae) has a

similar distribution as Dacrydium cupressinum and is
indeed abundant in the same type of forest, Podocarp-
mixed broadleaf forest from near sea level to ca. 700 m.
It is also much less common in South Island than in
North Island and in the former mainly stays near the
coast. Both genera have a palaeotropical Malesian-
Pacific distribution with extensions northward into
subtropical Indochina and southward to New Zealand
(map MA-43 on p. 316 and map MA-31 on p. 309). South
Island is at their climate limit, with Stewart Island at
47° S at the highest latitude for both genera (Dacry-
carpus reaches farthest north in Myanmar [Burma] at
nearly 27° N). This high latitude in the south is made possible by a strongly maritime climate
with little or no frost at low altitude near the coast. Podocarp-mixed broadleaf forest is often rich
in conifers, with Dacrydium cupressinum, Podocarpus totara, P. cunninghamii, Prumnopitys taxi-
folia, P. ferruginea, Halocarpus bidwillii, Manoao colensoi, and Phyllocladus trichomanoides join-
ing Dacrycarpus dacrydioides. In the far N of North Island Agathis australis can be added to this
mixture. However, these forests cannot be characterized as conifer forests and in this they differ
markedly from the diverse conifer forest of the Northern Hemisphere, such as occur in California,
Japan and elsewhere. Angiosperms are always an important part of the forest and succession, if
left undisturbed, would eventually oust nearly all of the conifers. It is the episodic setbacks of the
forest canopy by natural ‘disasters’ (or logging!) that will allow the conifers to persist indefinitely.
Only on unfavourable substrates, such as in peat bogs or on ridgetops, can conifers in New Zea-
land successfully compete with angiosperms. Dacrycarpus dacrydioides is a species that occurs in
all these habitats, but it can attain enduring dominance in lowland swamp areas.

Halocarpus

The genus Halocarpus (Podocarpaceae) is endemic to New Zealand. It is a sister clade to Para-
sitaxus (New Caledonia) and Lagarostrobos + Manoao (Tasmania and New Zealand) in a “Prum-
nopytoid” clade, a relatively basal clade in a phylogenetic analysis of the Podocarpaceae based on
molecular (DNA sequence) data by Biffin et al. in Turner & Cernusak (eds., 2011). The latter two
genera are monospecific, with Parasitaxus the only known parasitic gymnosperm. The divergence
time of this phylogenetic split is estimated to date back to the mid Cretaceous about 100 Ma. The
implication of this is that vicariance, rather than dispersal, best explains their current distribu-
tion, because the areas where these taxa occur were then still connected as a single landmass:
E Gondwana. In “vicariance biogeography” (Nelson & Platnick, 1981) populations are divided by
the emergence of geographical barriers, e.g. opening seaways, and the separate populations then
evolve into new species, leaving the parental species extinct. This pattern of evolution chimes
in with cladistic methodology and for some time dispersal events (which do not always fit the
cladistic paradigm) were anathema to cladists. Both explanations are possible and therefore valid,
and because we can never reconstruct the past with 100% confidence, a ‘best fit’ hypothesis is the
way forward to understand the biogeography of both extinct and extant species.
382 australasia

Map AU-59. Halocarpus biformis (black) [n=90, Map AU-60. Halocarpus bidwillii [n=81, m=81, h=4,
m=90, h=4, 1794–2006] and H. kirkii (red) [n=44, 1839–2005]
m=44, h=4, 1867–2007]

Halocarpus biformis and H. bidwillii are widely distributed and occur on all three main islands of
New Zealand, but are replaced by H. kirkii in the N of North Island. This distribution pattern sug-
gests that H. kirkii has ecological requirements different from the other two species. Its general
distribution coincides with that of Agathis australis and Libocedrus plumosa and, to a lesser
extent, with Phyllocladus toatoa. All are lowland forest conifers and the northern part of North
Island, reaching into the subtropical zone, has a very mild and equitable climate. Halocarpus
bidwillii and H. biformis are montane conifers, occurring between (400) 600 and 1500 m a.s.l. The
highest points in the N of North Island are around 700–900 m and much of the land is consider-
ably lower. It appears therefore that H. kirkii is limited by climate and can be considered as the
subtropical species in the genus. It is a tree to 25 m tall occurring in mixed conifer-angiosperm
forest. This species is listed as Vulnerable (VU) on the IUCN Red List. Halocarpus bidwillii is a
prostrate or spreading shrub, rarely a dwarfed tree, common in open tussock grassland or scru-
bland at higher altitudes in the N half of its range (both North and South Island). In Fiordland in
South Island it occurs with podocarp trees like Prumnopitys taxifolia, P. ferruginea and H. biformis
in open coniferous forest at lower altitudes. Halocarpus biformis is a small tree and often occurs
with or near the shrub-like species H. bidwillii but perhaps prefers a more forested habitat. In
open terrain it also becomes a dense shrub and when the two taxa grow side-by-side, they can
look very similar.

Halocarpus bidwillii in open grass-scrub

vegetation just above the forest edge in
Tongariro National Park, North Island,
New Zealand. The dark green trees in
the background are Phyllocladus tricho-
manoides. Halocarpus bidwillii is capable
of ‘layering’ whereby branches touching
the ground take root. In this way it forms
spreading bushes overcoming the difficul-
ties of competing with other plants as a
seedling. These large shrubs often remain
sterile for a considerable time, or produce
few cones at irregular intervals. Photo
credit Aljos Farjon.
 australasia 383

Manoao

Map AU-61. Manoao colensoi [n=86, m=85, h=5, 1841–2010]

Manoao colensoi (Podocarpaceae) is a tree with a seem-

ingly erratic distribution in North Island and a coastal
distribution in NW South Island. It occurs from near
sea level to ca. 950 m so it is not limited to mountains.
In Northland it is present from Cape Reinga to just
S of Hokianga Harbour and on the E coast at about
the same latitude; then there is a gap until it re­appears
in the hills just N of Auckland and, to the E, on Great
Barrier Island and the Coromandel Peninsula. Further
south, outside of Tongariro National Park, it has only
been found in a few locations in North Island. It is
abundant in Westland in South Island, but very rare
elsewhere. As a constituent of Podocarp-mixed broad­
leaf forest at lowland to sub-montane altitudes, it occurs with all the other conifers commonly
found in this forest type, but is apparently only common in certain areas. This species was for-
merly classified in Lagarostrobos (now a single species in Tasmania, see map AU-42 on p. 369)
but was separated on morphological distinctions, a taxonomy now confirmed by analysis of DNA
sequence data (Biffin et al. in Turner & Cernusak, eds. 2011). This makes it the second endemic
genus of conifers in New Zealand, with Halocarpus, to which it is related.

Lepidothamnus

The genus Lepidothamnus (Podocarpaceae) has three species, two occur in New Zealand and
one in S Argentina and S Chile. No fossil record is yet known of this genus, but evidently its spe-
cies must have occurred in other parts of Gondwana, in particular in Antarctica; this particular
disjunction pattern is known from several other southern conifer genera (e.g. maps AU-48,
AU-100, SA-5).

Map AU-62. Lepidothamnus intermedius [n=94, Map AU-63. Lepidothamnus laxifolius [n=82, m=81,

m=93, h=4, 1868–2006] h=4, 1839–2006]
384 australasia

Lepidothamnus intermedius is a shrub or small tree to 15 m tall, often multi-stemmed, and occurs
in swamp forests and other boggy places as well as on exposed ridges at altitudes from near sea
level to ca. 900 m. It is abundant on Stewart Island, quite common in Fiordland and in the NW
of South Island, and in Tongarira National Park and the Coromandel and Kaimai Ranges of North
Island. Lepidothamnus laxifolius is perhaps the most diminutive of all conifers. It forms slender,
creeping stems up to 1 m long, branching sparsely or sometimes more densely, rarely suberect and
usually trailing through the vegetation or forming low mats over rocks. It occurs in moorlands,
peat bogs, and tussock grass slopes in the mountains, usually between 750 and 1200 m a.s.l., on
Stewart Island down to sea level. In North Island it occurs in the higher mountains, but has not
been found on Mt. Taramaki (Mt. Egmont). In South Island it is absent from the highest, glaci-
ated part of the Southern Alps. These omissions are not easy to explain, but it is possible that
relatively recent geological events, such as volcanic eruptions and massive erosion caused by
expanding and retreating glaciers have played a part. Perhaps also this tiny conifer, while produc-
ing seeds subtended by a colourful, succulent receptacle, is not capable of long distance dispersal.
Re-colonizing lost ground would then be a very slow process taking thousands of years.

Phyllocladus

The genus Phyllocladus (Phyllocladaceae) has one of the most intriguing distributions of all coni-
fers. Its origin is similarly special, and to that topic we must pay attention first. The earliest fossil
records come from pollen found in Palaeocene strata in Australia and Patagonia (Morley in
Turner & Cernusak, eds. 2011). Phylogenetic analysis based on molecular (DNA sequence) data
and using ‘Bayesian relaxed-clock’ calculations estimated its origin to have taken place sometime
during the Late Cretaceous (Leslie et al., 2012). This and similar studies have placed Phyllocladus
in a basal clade but within a larger clade of Podocarpaceae; the taxonomic implication being that
if taxa = clades, this genus is a member of that family. This is controversial because morphological
and biological data, e.g. its unique phyllodes, an arillus that is not homologous to the epimatium
of Podocarpaceae, and aberrant pollination mechanisms indicate independence as a family, even
though it may have evolved from a primitive early member of Podocarpaceae (Farjon, 2010a). Its
origin is undoubtedly Gondwanan and of Mesozoic age and further fossil evidence is eagerly
awaited. Phylogenetic studies of the species of Phyllocladus report low resolution or weak statisti-
cal support of clades (Wagstaff, 2004), which could indicate recent origin but also slow mutation
rates. Below we shall see that in this case the latter is more likely.

Map AU-64. The distribution of the

genus Phyllocladus. Its notable absence
from the continent of Australia is
explained in the text.

Only four species are presently rec-

ognized, and they are distributed as
follows: Phyllocladus aspleniifolius in
Tasmania, P. trichomanoides (with
two varieties) and P. toatoa in New
Zealand, and P. hypophyllus in Male-
sia, from New Guinea to Borneo and
the Philippines. The great absence is
the continent of Australia, but
absence in New Caledonia is also
remarkable. Pollen records of Phyllo-
cladus are found in Australia from
the Palaeocene onwards through to
 australasia 385

the Holocene, so it became extinct there quite recently (Kershaw & McGlone in Enright & Hill,
eds. 1995). In New Guinea pollen fossils go back to the Miocene, but in Borneo they first appear
in the Pleistocene, and no records are known from further west on the Sunda Shelf (Morley in
Turner & Cernusak, eds., 2011). We can now reconstruct what is likely to have happened. The
genus Phyllocladus originated somewhere in Gondwana, possibly in what is now Antarctica, in
the Late Cretaceous. It spread to South America, where it became extinct during the Paleogene.
It also spread eastward to Australia and the peninsula that became New Zealand. In Australia it
could finally ‘hang on’ only in Tasmania, which is cool and wet. As Australia moved rapidly north-
ward in the Tertiary it passed into the high pressure climate zone of the Southern Hemisphere,
bringing desertification to all but the fringes, and from here Phyllocladus passed into New Guinea
in the Miocene but existed in the SE of the continent until perhaps only 1000 years ago. When
New Guinea accreted on to the Sunda Plate, through ‘island hopping’ the genus, without diversify-
ing into further species, travelled westward and northward, finally reaching W Borneo about
1.9 Ma.

Map AU-65. Phyllocladus toatoa [n=65, m=64, h=4,

1844–2010]

This species is limited to the northern half of North

Island, on or N of the 39° S parallel. It is a small tree
that tends to be rare or scattered in mature Kauri for-
est and Podocarp-mixed broadleaf forest, but becomes
more abundant on poorer sites where competition is
less intense. In secondary forest, which after a history
of logging is now much more common, it can establish
itself quickly as an opportunistic invader of more open
vegetation. It is difficult to say from the distribution
map to what extent this opportunism has influenced
its current extent of occurrence (EOO) as it may have
been present, but in lower numbers, in all these locali-
ties in the past.

Map AU-66. Phyllocladus trichomanoides var. trichoman-

oides [n=103, m=103, h=5, 1834–2010]

Phyllocladus trichomanoides var. trichomanoides is

more common than P. toatoa in the Kauri forests of
the northern parts of North Island. It attains greater
size in tall forest and can better compete in the can-
opy. Yet it has a similar role to play as an opportunis-
tic invader in the early successional stages after a fire
has destroyed the canopy. Larger trees of this species
can compete better and therefore stay longer as part
of the recurring Kauri (Agathis australis) dominated
forest. The ranges of these two species of Phyllocladus
overlap in the N of North Island, but P. trichomanoides
var. trichomanoides extends further S and into South
Island. In those localities it grows in montane mixed forest dominated by conifers such as Dacry-
carpus dacrydioides, Prumnopitys taxifolia and Manoao colensoi.
386 australasia

Map AU-67. Phyllocladus trichoman- Phyllocladus trichomanoides in Tongariro National Park, North

oides var. alpinus [n=100, m=96, h=4,
1839–1998]
Island, New Zealand. At this locality, the two varieties appear to
occur together, with var. alpinus represented by the shrubs in
the foreground. Nearly pure stands of relatively young trees indi-
cate an expansion of the forest, probably after volcanic activ-
ity of Mt. Ruapehu, an active volcano, destroyed earlier forest
cover. Photo credit Aljos Farjon.

Phyllocladus trichomanoides var. alpinus is the high montane to subalpine and usually shrubby
form of P. trichomanoides, in northern parts of its range occurring from 500 m to about 1800 m
but in the southern part descending nearly to sea level in exposed sites. It is the most widespread
and common variety in Southland, where in some localities it attains 8–9 m as a conical, densely
foliaged tree. In the ecotone between subalpine grassland and scrubland and forest, a mosaic of
open space and forest patches is often dominated by both varieties, with the tree form merging
into the forest, where other conifers and occasional angiosperms join in. In some accounts (e.g.
Eckenwalder, 2009; Debreczy & Rácz, 2011) this taxon is treated as a separate species.
 australasia 387

New Caledonia

Geography

New Caledonia is an archipelago situated in the SW Pacific Ocean about 1500 km E of Australia,
just N of the Tropic of Capricorn between ca. 19° 30′ and 22° 50′ S and 163° 30′ and 167° 40′ E in
the Coral Sea. It consists of a large, elongated and mountainous island, Grande Terre, ca. 420 km
long and 50–70 km wide, and several smaller islands, among which the flat Îles Loyauté to the
E are the largest and the Île des Pins at the SE end of Grande Terre is a small mountain. All the
islands are surrounded by coral reefs, of which the Grand Récif around Grande Terre is the second
largest in the world, after the Great Barrier Reef. The highest mountains are Mt. Panié (1628 m)
in the north and Mt. Humboldt (1618 m) in the south. The steep E coast, where Mt. Panié rises
directly from the ocean, receives much of the rainfall (more than 8 m on this mountain every
year) whereas the W coast, in the rain shadow of the mountains, is relatively dry. Temperatures
are nearly constant throughout the year, moderated by the influence of SE trade winds, but at
22–24° C near sea level tropical nevertheless. However, the islands lie in the path of cyclones,
which are most frequent from November to March. The coral reefs protect the coasts from the
ocean surf, trees grow up to the water line and mangroves are common at the outlets of rivers.
The complex geology of Grande Terre is one of the most influential physical features relevant to
plant growth and to conifers in particular. Ancient Palaeozoic rocks form the core of the island,
but they surface mainly in the N, culminating in Mt. Panié. These are acidic or neutral substrates
for plants. Overlying these formations elsewhere is a complex array of rocks of volcanic or sedi-
mentary origin, the most interesting of which are metamorphosed volcanic rocks rich in metals,
collectively known as ultramafic rock. These can be bedrock or reworked materials, and are rich
in iron, manganese, nickel, chrome, cobalt and other metals. They are poor or deficient in miner-
als such as nitrogen, calcium and phosphor, needed for plant growth. Such substrates are ubiq-
uitous in the southern part of the island, but also present on the massifs across the length of the
island and on the highest point of the Île des Pins. They are not found on the Îles Loyauté, which
are uplifted coral atolls. These ultramafic rocks play a major part in the distribution and diversity
of conifers in New Caledonia. The flora is very rich, with over 3000 species so far recognized, 80%
of which are endemic, indicating a long evolution during isolation.

Conifers in New Caledonia

Families Genera Species Species + infra Countries with Area of occupancy Endemic taxa
conifers in km²
4/8 14/70 43/615 43/794 1/1 1845/18,575 43
Numbers after ‘/’ are global or country totals.

Given its small area, New Caledonia is the most diverse conifer territory in the world, with 43
species, 14 genera (three endemic, one shared only with Fiji) and four families present. For the
genus Araucaria (Araucariaceae) it is the world centre of diversity, with 13 out of a world total
of 19 species. Nearly half of all species in the family Araucariaceae occur here, as there are also
four species of Agathis present. There are some very unusual conifers too, with Austrotaxus (Tax-
aceae) a monospecific genus and only representative of this family in the Southern Hemisphere,
and Parasitaxus (Podocarpaceae), another monospecific genus and the only parasitic gymno-
sperm known in the world. Neocallitropsis (Cupressaceae) is the third monospecific and endemic
genus in New Caledonia. Retrophyllum minus (Podocarpaceae) is the only true rheophyte among
gymno­sperms. There are rare species confined to very small areas or even limited to a few hun-
dred mature plants, and common species that occur all over Grande Terre. Perhaps most remark-
388 australasia

able, none of the 43 species occur elsewhere, conifer endemism in New Caledonia is 100%. It
is entirely appropriate to treat New Caledonia as a distinct region in Australasia in this Atlas,
despite its diminutive size compared to Australia or even New Zealand. Due to its small size, it is
not very informative to give a map showing all species together, as has been done for the larger
areas in this chapter. Conifers would appear to occur nearly everywhere. Instead we present a
map on diversity and maps for some genera and for all species. Maps of species have been pre-
sented before in Vol. 4 of the Flore de la Nouvelle Calédonie et Dépendances (De Laubenfels, 1972),
similarly based on herbarium records. Forty years later, not only have more specimens been col-
lected, but access to these resources has been greatly improved. As a result, our maps are more
complete, with many new localities added since that first effort to map the distribution of New
Caledonia’s remarkable conifer diversity.

Map AU-68. Number of species in 6 minute cells with

two cells located on Montagne des Sources/Rivière
Bleue with 19–23 species. Province Sud in general has
greater conifer diversity than Province Nord. The ultra-
mafic massifs and especially the ‘Massif du Sud’ have
the greatest diversity. This emphasises the specializa-
tion of many conifer species in New Caledonia in cop-
ing with these more or less toxic soils. Areas without
conifers are mostly situated on the W coast, in the rain
shadow of the mountains, and in the mountainous
�‒�
�‒�
parts of Province Nord on acidic or neutral substrates,
��‒�� where competition from angiosperms in the rainforest
��‒�� may exclude conifers.
��‒��

Map AU-69. Geological map

of New Caledonia. The ophi-
olitic nappe indicated in black
forms the ultramafic massifs;
the large area of this forma-
tion in the southern part of
Grande Terre is known as the
Massif du Sud. Source: Journal
of Pet­­rology 53 (5), May 2012.
© Oxford University Press.

Araucariaceae

The family Araucariaceae has three genera, two of which are present in New Caledonia. The
genus Agathis has a Malesian-Pacific distribution, and although it is represented in New Caledo-
nia with four species out of a total of 17 species, the majority occur in Malesia and the genus is
therefore discussed in that chapter (map MA-66 on p. 332). Here we present maps for the species
in New Caledonia. The genus Araucaria has 13 species in New Caledonia and the island is its
centre of diversity.
 australasia 389

Agathis

Map AU-70. Agathis moorei [n=76, m=56,

h=9, 1861–2002]

Agathis moorei (syn. A. corbassonii) is

widespread in the rainforests of Grande
Terre. It occurs as solitary trees or small,
dense stands in angiosperm-dominated,
dense forest at altitudes from 30 m to
700 or even 1000 m. In the south of the
island it is found near the coast on the
W side, but where the rain shadow
effect from higher mountains becomes
apparent, it retreats into the interior or
moves over to the E coast. This species
can grow on soils derived from a variety
of usually metamorphosed rock types,
but avoids ultramafic substrates. As most of the large rainforest trees do the same, it has to com-
pete with these; presumably it takes advantage of gap formation from fallen trees, probably
mainly larger gaps created by cyclones or landslides, as is suggested by the occurrence of small
stands or groves of this species. Foresters in New Caledonia noted that there are colour differ-
ences in the barks of trees and assigned those with a reddish bark to A. corbassonii. It appears
that this is not a good taxonomic character but is influenced by exposure to light, which turns
the bark grey as in the traditional concept of A. moorei (Farjon, 2010a, pp. 161–162). Heavy exploi-
tation has made large trees scarce in many areas (and exposed the bark of remaining trees to
sunlight!) and it is possible that this has had an impact on the distribution pattern shown on this
map (but see the next species for comparison). Agathis moorei is listed as Vulnerable (VU) on the
IUCN Red List.

Map AU-71. Agathis lanceolata [n=161,

m=140, h=7, 1870–2003]

Agathis lanceolata is common in the

southern part of Grande Terre, occur-
ring in the patched remains of rainfor-
est in the hills and low mountains of
the ‘Massif du Sud’, the largest extent of
ultramafic rock and soil on the island. It
is more scattered further north, where
it is found in forested valley bottoms in
more remote locations. On Mt. Dzumac
it reaches its highest altitude between
700–900 m, but in the ‘Massif du Sud’ it
can be found as low as 50 m and usually
not higher than 500 m. This species is
almost exclusively growing on ultramafic red soil, an erosion product from peridotite and other
metal-rich base rocks with high iron content. It can become a large tree to 50 m tall, so logging
has been extensive and few of these large trees can be found today. As with other species in the
genus, it becomes an emergent in mixed rainforest, usually solitary or a few trees together, but
in a younger phase after regeneration small stands of many trees can occur in former forest gaps.
390 australasia

The rarity of A. lanceolata beyond the southern, lower part of Grande Terre is unlikely to be the
result of logging, as this has certainly been as intensive, or more so, in the ‘Massif du Sud’, closer
to Nouméa and easier to construct forest roads in. The distribution pattern, based on extensive
collecting by Timothy Waters (University of Oxford) and others, is likely to reflect the natural
situation. Agathis lanceolata is listed as Vulnerable (VU) on the IUCN Red List.

Map AU-72. Agathis ovata [n=152, m=139, h=10, 1861–2003] Agathis ovata on a low mountain near Yaté,
New Caledonia. This species occurs mostly in ‘maquis minier’ vegetation on ultramafic substrates and there
it remains a small, flat-topped tree. Photo credit Aljos Farjon.

Even more so than the previous species, Agathis ovata is limited to the southern part of Grande
Terre; just two localities are known from the central part of the island. This species is not a rain-
forest emergent, but occurs in lower vegetation known locally as ‘maquis minier’. This type of
vegetation reminded the French of the maquis along the Mediterranean coast, but of course its
floristic composition is entirely different. The reference to mining alludes to the metal-rich ultra-
mafic rock to which this species (and the maquis) is mostly confined. In a few places it also
penetrates into the forest ecotone, but as it is a small tree it cannot compete with tall rainforest
trees so it stays in the margins. Its altitude range is substantial, from 30 m to 1050 m, and it
extends on to the higher ridges of the Montagne des Sources, Mt. Dzumac and other southern
mountain massifs. Here it may be associated with Araucaria and Nothofagus and, due to competi-
tion with these taller trees grow to 20–25 m tall. Not logging, but fire is the main threat to this
species, as it occurs in vegetation as prone to burning as its Mediterranean namesake. Agathis
ovata is listed as Endangered (EN) on the IUCN Red List.

Map AU-73. Agathis montana [n=37, m=37, h=4,

1950–2003]

Agathis montana is restricted to the Mas-

sif du Panié (Mt. Colnett, Mt. Ignambi, Mt.
Panié) and the Roches d’Ouaième in the NE of
Grande Terre. On these mountains it occurs in
high montane forest between 950 m (Roches
d’Ouaième) and 1050–1600 m with Retrophyl-
lum comptonii (map AU-101 on p. 410), the only
other large tree. The understory is made up of
shrubs and many ferns. The crowns of A. mon-
tana spread very wide in candelabra fashion
and support mossy and ferny epiphytes. The
Massif du Panié receives the highest rainfall
in New Caledonia, in excess of 8 m per annum
 australasia 391

and water is tumbling down the steep E slopes in numerous never drying waterfalls. Cloud is
nearly always covering the mountain and the summit ridges where this species is most abundant
are shrouded in fog almost year-round. The massif is composed of ancient micaschist with a low
pH and the runoff is slightly acidic, in contrast to the ultramafic rock on which most New Cale-
donian conifers occur.

Araucaria

The genus Araucaria has 19 species, 13 of which occur in New Caledonia. These belong to Sec-
tion Eutacta, of which the only other two species are A. cunninghamii (Australia, New Guinea,
map AU-10) and A. heterophylla (Norfolk Island, map AU-118). The remaining four species belong
to three sections: Section Araucaria with A. araucana and A. angustifolia (South America, maps
SA-30 on p. 449 and SA-23 on p. 443), Section Bunya with A. bidwillii (Australia, map AU-9) and
Section Intermedia with A. hunsteinii (New Guinea, map MA-65 on p. 331). Their distribution is
highly disjunct, a relict of a nearly world-wide distribution that dates back to the Jurassic and
included both Gondwana and Laurasia. When these two supercontinents broke up, Araucaria
became successively extinct in Eurasia, North America, Africa, India and Antarctica and was
greatly reduced in Australia and South America. This reduction is reflected in the taxonomy,
supported by phylogenetic analysis based on DNA sequence data; three of the four sections have
1–2 species each left of what must have been far greater diversity. The higher species number in
Section Eutacta is entirely due to the diversity of the genus in New Caledonia. In South America,
the two species occur far apart on two sides of the southern half of the continent, in warm tem-
perate to cool temperate latitudes. In Australasia (including New Guinea) the remainder of the
species are limited to the eastern fringe of Australia, New Guinea, New Caledonia and Norfolk
Island. These areas are mostly tropical but in Australia extend southward to warm temperate lati-
tudes. Absence is conspicuous in SE Australia including Tasmania and in New Zealand. In these
locations, as elsewhere, the genus became extinct.

Araucaria
Eutacta
Bunya
Intermedia

Map AU-74. The distribution of the genus Araucaria (19 species divided into four sections) is highly dis-
junct and limited to South America and Australasia (including New Guinea).

The diversity of species in New Caledonia is most likely of geologically recent origin. Phylogenetic
analysis has shown that these species are closely related, even though morphologically some are
highly distinct. The sister species to all the New Caledonian species is likely A. heterophylla from
Norfolk Island, and that clade is again related to A. cunninghamii (Setoguchi et al., 1998) as the
basal species in Section Eutacta. The relationships among the 13 New Caledonian species remain
poorly resolved (Gaudeul et al., 2012); apparently the plastid gene and AFLP sequences studied
show little variation among these species. From these results it is inferred that a single ancestor
of these species gave rise to the diversity and that this radiation occurred while New Caledonia
was an island separated from Australia. How long ago this speciation started is very difficult to
392 australasia

estimate in the absence of a fully resolved phylogeny. New Caledonia separated from Australia ca.
80 Ma so there seems to have been ample time, but on the other hand the speciation was prob-
ably linked to geological upheavals which occurred mostly during the Eocene, creating the differ-
ences in altitude, rock and soil type and precipitation that created different habitats. It has been
proposed on geological evidence that the entire island has been submerged for several million
years during this epoch and that therefore the current species of plants and animals all evolved
from ancestors that arrived after 37 Ma when New Caledonia was a remote island. While this is
possible for some bird dispersed conifers (mainly Podocarpaceae, see Enright & Jaffré in Turner &
Cernusak, eds. 2011) it is highly unlikely for Araucariaceae, which are not known to be dispersed by
birds and of which species the seeds only travel short distances. Of the 13 species of Araucaria in
New Caledonia, only three also occur on the satellite islands, all the others are confined to Grande
Terre. Nine are virtually restricted to ultramafic rock or soils, two are ­indifferent, ­occurring on
these as well as other substrates, and one, A. schmidii is entirely absent from ultramafic soils (see
Table 5 on p. 398). Araucaria columnaris grows predominantly on raised coral reefs but can be
found near the coast on basalt or serpentine. This is also the species that occurs on the smaller
islands, as well as on the coast of Grande Terre. The distribution of the genus Araucaria in New
Caledonia therefore coincides to a large extent with the presence on or near the surface of ultra-
mafic rock (Farjon, 2008, p. 268–271). This puts the trees into a conflict situation with the open
pit mining for nickel, an activity that poses the greatest threat to several species of Araucaria with
limited distribution and/or small populations.

Map AU-75. The distribution of the genus Araucaria in New Caledonia. This map excludes evident plant-
ings of especially A. columnaris, a popular tree with the Melanesian first arrivals, and some attempts at
forestry using several species by Europeans in the southern part of Grande Terre. Some dots may represent
populations now extinct due to nickel mining.
 australasia 393

Map AU-76. Araucaria columnaris [n=67,

m=54, h=14, 1774–2005]

Araucaria columnaris is the iconic spe-

cies of the genus in New Caledonia. It
is the species first seen on approaching
the SE coast of Grande Terre, as James
Cook famously did when he ‘discov-
ered’ and named New Caledonia during
his second circumnavigation in 1774.
It is also the only species native to the
Îles Loyauté and Île des Pins. This spe-
cies was and still is popular with the
Kanaks, the Melanesian first arrivals on
the islands, and they have planted it on
the Îles Loyauté, on the coast of Grande
Terre and in valleys in the interior of the
main island. The map includes none of these and is an attempt to represent natural distribution,
not an easy task in some cases. The location at Baie des Tortues (Turtle Bay) about halfway up the
W coast (red circle) is a case in point. Here trees grow on low slopes around a bay where turtles
used to bury their eggs in the beach, a popular gathering spot for the Kanaks. Did they plant
the trees? The population looks natural, with uneven aged trees and some regeneration ­present.
Although often confined to raised coral reefs, this species does grow on low slopes of basalt or
serpentine, e.g. at Port Boisé and on rock slopes just S of Goro. But Baie des Tortues is an oddly
out­lying location, especially as it is on the sheltered W coast. Although trees are toppled, A. colum-
naris is able to withstand the force of cyclones, which tend to hit New Caledonia from the SE,
exactly where it is naturally distributed. Individual trees grow 50 m tall but are very columnar
indeed; branches are ripped off in a heavy storm, leaving not much more than a standing mast,
and regrow from dormant buds. It seems the perfect adaptation to a stormy coast.

Araucaria columnaris on raised coral reefs at the Baie d’Upi, on the Isle of Pines. Their narrow columnar
crowns resist cyclone winds. Photo credit Timothy Waters.
394 australasia

Map AU-77. Araucaria schmidii [n=16, m=16, h=8,

1951–2005]

Araucaria schmidii is the only species that does

not grow on ultramafic rock. It is only known
from the summit of Mt. Panié in the NE of
Grande Terre, which is composed of acidic mic-
aschist, a metamorphosed igneous rock type.
This is the rarest species in the genus, with a
population of only a few hundred trees. Explo-
rations along the chain of the Massif du Panié
by the late ecologist and conservationist Henri
Blaffart and local guides so far failed to find any
other populations of A. schmidii (pers. comm.
to AF, Nov 2005) although Nasi (1982, p. 64)
recorded finding it on nearby Mt. Colnett. The
other species present at lower altitude is
A. montana; perhaps there was some confusion
with this common species. Araucaria schmidii grows mostly on the rocky edges of the summit
plateau, at 1400–1630 m, as the only tree or with Agathis montana. On the IUCN Red List it is
classified as Vulnerable (VU) based on its small population only, susceptible at least in theory to
‘stochastic’ events. The location is very remote and can only be visited by the most determined
bush hikers. The forest on Mt. Panié is within a protected area, proposed as part of a World Heri-
tage Site.

Map AU-78. Araucaria montana

[n=80, m=70, h=14, 1869–2005]

Araucaria montana is the most

widespread and common species
of the genus in New Caledonia. It
is indifferent to rock type and its
limitation is low altitude; not
found below 200 m it usually
occurs between 800 and 1200 m
and up to 1400 m. In part, this
explains its rarity in the S of
Grande Terre, although the Mon-
tagne des Sources and Mt. Dzou-
mac (above 1000 m) where it is
absent would appear to be suit-
able habitat. The one location in
the south on the map is on the
Rivière Bleue at 600–800 m, based on a collection by D. J. de Laubenfels kept at Paris (P) but not
confirmed by A. Farjon and R. Mill during their visit to Paris in October 2005. It is perhaps of
uncertain identity. Araucaria montana is commonly found in low forest or tall maquis on ridges
and in taller forest in ravines, where it tends to occupy the steeper rocky sides and slopes. As with
 australasia 395

several other species, it is an emergent rising well above the general canopy, occasionally reach-
ing 40 m tall but usually considerably smaller. This species is listed as Vulnerable (VU) on the
IUCN Red List.
Map AU-79. Araucaria subulata [n=44, m=27, h=10,
1861–2005]

Araucaria subulata occurs mainly in the south-

ern part of Grande Terre (‘Massif du Sud’ of
geologists) and in a few localities in the cen-
tral mountains. One of these is the Table Unio,
the plateau from where this species was first
described. Its altitude ranges from 150 to 1070
m and it is a species restricted to ultramafic
rocks and soils. Araucaria subulata is a tall,
columnar emergent tree in dense tropical
rainforest with a canopy of 12–30 m, above
which it can rise 20 or more metres. It is com-
mon in steep valleys and ravines on the rain-
ward side (E and SE) of mountains. It can occur with another emergent conifer, Agathis
lanceolata (map AU-71 on p. 389 and both would need gap openings of some considerable size,
if not larger removal of the canopy, to regenerate successfully. The natural disturbing agents
would be cyclones or fires. The distribution of both conifers on the island may be related to this
mode of perpetuating these species in the forest. Increased frequency of fires under human
influence is working against successful regeneration.

Map AU-80. Araucaria bernieri [n=49, m=40, h=13,

1942–2005]

Araucaria bernieri has a similar distribution to

A. subulata but is somewhat more common; it
also has a similar ecology. Indeed, sometimes
the two species occur together in the same for-
est. The altitude range of A. bernieri is some-
what lower, commonly at 100–500 m and rarely
to 800 m, so this is the commonest species in
forests on the low mountains of the far SE of
Grande Terre. This species has been observed
to follow stream beds in ravines; perhaps these
wetter places form some protection against
fires. It is always limited to ultramafic sub-
strates, on which the forest canopy tends to be
lower and often more open. Both A. bernieri and A. subulata can attain 50 m in dense forest.
Decline of the total population is inferred from continuing destruction or disturbance of its forest
habitat, with only some of its known localities in protected areas. It is listed as Vulnerable (VU)
on the IUCN Red List.
396 australasia

Araucaria bernieri on Pic Grand Kaori, New Caledonia. Tall spires of these trees are emerging well above
the canopy of much lower angiosperms (and a few conifers such as podocarps). The typically solitary trees
are probably the long time survivors of a regeneration phase after episodic forest disturbance by a cyclone
or a fire. Photo credit Aljos Farjon.

Map AU-81. Araucaria biramulata

[n=33, m=31, h=13, 1864–2005]

Araucaria biramulata is an
uncommon species with a scat-
tered distribution across Grande
Terre, limited to ultramafic sites.
Most of the localities where her-
barium collections were made are
in Province Sud, but this species
is also known from two localities
in Province Nord: Mt. Kaala (far-
thest N) and Massif de Kopéto.
Both localities have large active
nickel mines, and in neither have
new collections been made since
1969. On Mt. Kaala, this species
was collected in 1943 and 1968; a
day long search by AF in 2005 remained without results. That mountain’s upper slopes and sum-
mits or ridges have almost entirely disappeared, together with all vegetation, and only A. montana
was seen there on what was left of the upper parts of the mountain (Farjon, 2008, p. 267). The
collection from the Massif de Kopéto dates from 1969, but there more forest remains and it could
still be present. No other species of Araucaria have been collected there as far as we know. Arau-
caria biramulata is listed as Vulnerable (VU) on the IUCN Red List; in two of its known localities
it may be extinct.
 australasia 397

Map AU-82. Araucaria laubenfelsii

[n=56, m=47, h=13, 1947–2005]

Araucaria laubenfelsii has its

main range in Province Sud, but
is known from three localities in
Province Nord, near Boakaine,
on Mt. Kaala and on Île Art, a
small island 50 km NW of the
tip of Grande Terre. It has prob-
ably disappeared from Mt. Kaala,
from where it was last collected
in 1968. In the south, it is abun-
dant in mountains N of Noumea:
Mt. Do, Mt. Dzumac, Mt. Mou,
Montagne des Sources and other
localities. These are mostly within
protected areas and are not sub-
ject to mining. Araucaria laubenfelsii has been found in different habitats ranging from maquis
to montane rainforest, but nearly always on ultramafic rock or soil.

Map AU-83. Araucaria luxurians. [n=51, m=45, Araucaria luxurians on the shore of Baie des Pirogues,
h=13, 1867–2005] Province Sud, New Caledonia. Photo credit Aljos Farjon.

Araucaria luxurians has a scattered distribution, ranging from Baie du Prony in the far south of
Grande Terre to the Îles Bélep 50 km NW of the main island. On Mt. Kaala, the northernmost
point in Grande Terre, it may be another victim of nickle mining; its most recent herbarium col-
lection was made in 1967 and it was not found during a search in 2005 (Farjon, 2008, p. 267). This
too is a species occurring on the ultramafic substrates. Araucaria luxurians is mostly found near
the coast, where it has some of its largest sub-populations, and is less common in the interior
mountains. This species is forming escarpment forest on serpentine cliffs next to the sea or in
the interior, where it can occur to 1000 m. It sometimes grows near stands of A. columnaris on
the coast in the south, but does not join it on the raised coral reefs. Considerable decline mostly
due to mining and a lack of protected areas that include this species have caused it to be listed
as Endangered (EN) on the IUCN Red List.
398 australasia

Map AU-84. Araucaria scopulorum

[n=55, m=54, h=10, 1943–2005]

Araucaria scopulorum is almost

entirely a coastal species, often occu-
pying steep slopes above the coral
lagoons from just above the beach to
the crest of the escarpment at 400–
600 m a.s.l. Its main range is along
the central E coast of Grande Terre,
between Thio and Ponérihouen, a
smaller area is on the NW coast, at
Mt. Poum and Dôme de Tiébaghi,
both of which are mined and have
no formal protection. These two
localities have been recently identi-
fied by A. Farjon and R. Mill from
previously misidentified herbarium specimens in Paris (P) and elsewhere. There is still a regen-
erating population at the southern end of Mt. Poum (AF pers. obs. Nov 2005). The distribution of
this species nearly coincides with nickel mining areas and many populations are in or adjacent to
active nickel mines. Whereas the number of known localities may actually have increased since
De Laubenfels (1972) published a map based on similar data, the number of trees has undoubt-
edly decreased due to widespread mining and lack of formal protection. This species is listed as
Endangered (EN) on the IUCN Red List.

Table 5. The species of Araucaria in New Caledonia and the three types of substrate on which they occur
naturally. Nine are limited to ultramafics, one is more common on raised coral reefs, two also occur on
acidic or neutral rocks or soils, and one is found on acidic rock and not on ultramafics.
 australasia 399

Map AU-85. Araucaria rulei [n=65, m=54, h=14,

1858–2005]
Araucaria rulei SE of Baie de Poro
(Néjeré Dumwà) a heavily disturbed
area with erosion caused by pros-
pecting and mining for nickel. Photo
credit Aljos Farjon.

Araucaria rulei is most common in the southern half of Grande Terre but there are a few scattered
locations in the northern half of the island. The isolated location on the Dôme de Tiébaghi in the
far north of the main island was discovered in 1975. This species is predominant in ‘maquis minier’
on ultramafic substrates and only rarely occurs in low forest. The altitude range is substantial, from
150 to 1150 m, but mostly it occurs between 400–800 m a.s.l. It is a sparsely branched, modest size
tree, usually not exceeding 20 m tall. Although sometimes forming populations several km² in
extent, the trees are scattered and do not form a forest. Despite its fairly wide distribution, A. rulei
is intricately linked with the destructive nickle mining and if not destroyed outright, its habitat is
often degraded by erosion from vehicle tracks, increased incidence of fire that comes with ease of
access and deliberate drainage ‘improvement’ to allow movement of heavy vehicles. Again, few
sub-populations are within protected areas and decline is ongoing, with large numbers of trees
dying from these adverse conditions. Araucaria rulei is listed as Endangered (EN) on the IUCN
Red List.

Map AU-86. Araucaria muelleri [n=49, m=41, h=14,

1868–2005]

Araucaria muelleri is in some respects similar to

A. rulei. It is an even more sparsely branched tree
with larger foliage branches and larger leaves,
similarly appearing as terminal tufts on primary
branches. Its ecology is also similar, as it occurs
most often in maquis, but sometimes in dense
forest with Nothofagus forming the lower canopy.
Its distribution is different, because it is restricted
to the ultramafic ‘Massif du Sud’, which is not a
mountain massif but a complex terrain of plains,
valleys with lowland rivers and some lakes (includ-
ing a large man-made reservoir) and hills and low
400 australasia

mountains. The ultramafic rock of the lowland plains and hills has weathered into laterite capped
by a hard, brown layer of ironstone, into which creeks and rivers have cut gullies. In such terrain,
sparsely vegetated, A. muelleri can be the tallest tree even though it rarely exceeds 15 m. The
ironstone/laterite is ultramafic and contains nickel, but in lower concentrations than the garni-
erite in the mountains; for this reason mining activity has so far been limited to exploration
surveys. However, much of the area is under mining concessions and very little of the land is in
nature reserves. At present, fires are the main threat to the small populations, often limited to
fewer than 20 or 10 trees, while regeneration is sporadic and slow. This species is listed as Endan-
gered (EN) on the IUCN Red List.

Map AU-87. Araucaria humboldtensis [n=31, m=29,

h=13, 1938–2005]

Araucaria humboldtensis has a limited distribu-

tion in the S of Grande Terre, occurring on the
highest mountains there, including Mt. Hum-
boldt. Other mountains with populations of
A. humboldtensis are Mt. Mou and the Montagne
des Sources. The altitude range is 750–1600 m
and at the highest point this species nearly
reaches the summit of Mt. Humboldt. Trees
are small to moderately tall (20 m) depending
on the vegetation they grow in, which is elfin
forest near the summits and on the highest
ridges; yet they are invariably emergents over
the general canopy. The substrate is ultramafic,
derived from serpentine or peridotite, but the content of nickel is often lower than elsewhere and
mining is not an issue of great concern for this species. Forest and bush fires are the main threat
to A. humboldtensis, increasingly set off by careless people. This species is listed as Endangered
(EN) on the IUCN Red List.

Map AU-88. Araucaria nemorosa Araucaria nemorosa at Port Boisé, New Caledonia, a Critically
[n=34, m=32, h=11, 1968–2005] Endangered species. Photo credit Aljos Farjon.

Araucaria nemorosa is a rare species with a very limited distribution. It is known from three
localities in the far S of Grande Terre. The largest and best known population is at Port Boisé;
another, much smaller one is situated a few km E of this on the coast at Cap Reine Charlotte, and
 australasia 401

a third at a place called Forêt du Nord (black circle), where only a few trees have been seen by
Timothy Waters in 2001. The main ­population is on a more or less level plain 20–40 m above the
bay of Port Boisé in low open woodland or tall maquis interspersed with open rocky ferritic soil
devoid of plant growth (‘cuirasse de fer’). The araucarias form small stands or grow solitary, over
not more than 2 km² in about eight sub-populations. Apart from mining on the fringes (so far)
there is the hazard of increased fires from both tourists and natives. The land is not a protected
area and in private ownership but with free and easy access to all. This species is listed as Criti-
cally Endangered (CR) on the IUCN Red List.

Cupressaceae

The Cupressaceae are represented in New Caledonia with three genera and six species. The taxo-
nomic relationships are with Australia and New Zealand. Libocedrus has three species in New
Caledonia and two species in New Zealand, Callitris has two species in New Caledonia and 13
species in Australia (map AU-16 on p. 354) and Neocallitropsis is a monospecific genus endemic
to New Caledonia, but related to Actinostrobus and Callitris. Libocedrus is known from the fossil
record of the Paleogene in Australia and Tasmania. These relationships may reflect the land con-
nections that existed between these islands and Australia until they were separated in the Late
Cretaceous and the Tasman Sea opened up between them. The fossils known at present have to
be interpreted as minimum ages; it is more significant to know that this paleo-geographic link
existed and older fossils may well be found in future.

Libocedrus

Map AU-89. Libocedrus austrocaledonica

[n=29, m=29, h=5, 1868–2005]

Libocedrus austrocaledonica is the more

common of the three species that occur
in New Caledonia, although its occur-
rence is concentrated in the mountains
of the southern part of Grande Terre.
Only one location, Mont Paéoua is
known in Province Nord. It is a shrub
or small tree growing among other coni-
fers and angiosperms on high mountain
ridges between 750 and 1400 m. On these
ridges the forest remains sparse and
low, with the occasional emergent tree
of Araucaria humboldtensis as excep-
tion. Competition from larger trees is
excluded, and characteristically, shrubs and trees on these nutrient poor and exposed ridge crests
have small leaves, allowing more light to filter through to the ground. If plants can cope with the
nutrient deficiency, as many conifers can, their seedlings can grow up here. Just a few metres
downslope soils are deeper, trees are larger with bigger leaves, and seedlings of most conifers
are denied sufficient light to grow. Such limitations cause rarity because in terms of surface area
these mountain ridges take up a tiny part of the topography. In the case of L. austrocaledonica a
further limitation is that it seems to avoid ultramafic rocks and is only found on humic, strongly
acidic substrates.
402 australasia

Map AU-90. Libocedrus chevalieri (black) [n=12, m=12,

h=3, 1940–1985]; L. yateensis (red) [n=23, m=23, h=5,
1947–2005]

Libocedrus yateensis in dense riparian scrub

along the Rivière Bleue, New Caledonia. Photo
credit M. Berenbrinker.

These species are very rare, each is known from only three locations. Libocedrus chevalieri has
been collected on Mt. Humboldt and Mt. Kouakoué in Province Sud and on the Massif de Ton-
Non on the E coast near Poindimié in Province Nord. Libocedrus yateensis occurs in a few locali-
ties along the Rivière Bleue-Yaté River and the Ouinné River in the far south and near Povila
in Province Nord. Libocedrus yateensis is a riparian species growing on the banks of creeks and
rivers at lower altitudes from 150 to 600 m, occupying river terraces and sometimes forming small
groves. Libocedrus chevalieri is a montane species growing on steep slopes in the contact zone
between schists and serpentines at altitudes between 650 and 1618 m (the latter is the summit of
Mt. Humboldt). Both species have very small populations, and are in decline from various causes.
Libocedrus chevalieri is listed as Critically Endangered (CR) and L. yateensis as Endangered (EN),
both up a category from the last assessments in 1998, on the IUCN Red List.

Callitris

Map AU-91. Callitris sulcata [n=29, m= 19, h=7, 1862–1983]

Callitris sulcata is a rare species occurring in the valleys

of the Comboui, Dumbea and Tontouta Rivers and
some tributaries in Province Sud on Grande Terre. It
often follows rivers and creeks where it grows on ser-
pentine slopes above the streams, but can also be
found on ultramafic soils in ‘maquis minier’ or on the
edges of gallery forests along rivers. It is a lowland spe-
cies with an altitude range between 40 and 200 m. The
shrubs or small trees can be difficult to find in dense
vegetation, unless they form thickets with several
 australasia 403

i­ ndividuals together. Some populations are within protected areas but others are not and may be
subject to logging or other forms of forest clearing even though this small tree is not a target spe-
cies. Increased fires are also threatening this species, which is listed as Endangered (EN) on the
IUCN Red List.

Map AU-92. Callitris neocaledonica [n=28, m=28, h=6,

1902–1988]

Callitris neocaledonica is also limited to the southern-

most part of Grande Terre, but it occurs at higher alti-
tudes than C. sulcata, from 560 to 1500 m. It is restricted
to ultramafic rocks and grows on slopes or along creeks
in the hills and mountains of the ‘Massif du Sud’, where
these metal-enriched rocks and soils are nearly con-
tinuous throughout the area. It is protected in several
reserves and, so far, has not been affected by mining
activities. Its natural habitat is ‘maquis minier’ or low
sclerophyll forest which is not subject to logging or
forest clearing.

Neocallitropsis

Neocallitropsis pancheri on the Rivière des Lacs, Plaine des Lacs, New Caledonia. It forms thickets in ‘maquis
minier’ on ultramafic ‘cuirasse de fer’ in this area. Photo credit Aljos Farjon.
404 australasia

Map AU-93. Neocallitropsis pancheri [n=37,

m=34, h=7, 1861–1996]

Neocallitropsis pancheri is one of the

more peculiar conifers of New Caledo-
nia. It is the only species of its genus,
of which no records, fossil or living, are
known from elsewhere. The genus is
related to Actinostrobus and Callitris or
to a more inclusive clade of ‘callitroid’
genera and could perhaps represent a
more primitive ancestral group of species
in a tribe Callitrideae that would include
fossil taxa. Its distribution in New Cale-
donia is almost restricted to the Plaine
des Lacs and surrounding hills between
Yaté in the north and Prony in the south,
but there are two outlying populations known. One is in the Montagne des Sources near Pic Buse,
the other locality is Mt. Paéoua in Province Nord (black circle). Both are higher altitude localities,
from 700 to 820 m in the Montagne des Sources to 1140 m on Mt. Paéoua. In the Plaine des Lacs it
occurs from 30 to 260 m a.s.l. and is much more abundant, often forming thickets. There it often
follows streams, but stays above any flooding except in extreme circumstances. The inundation
by the Yaté dam in 1959 is covering a vast area (in red laterite mud at low water levels) and must
have destroyed many sub-populations. This species is often associated along streams with the
podocarps Dacrydium araucarioides, D. guillauminii, and Dacrycarpus vieillardii, elsewhere with
Agathis ovata, Callitris neocaledonica, Podocarpus novae-caledoniae, and angiosperms. Despite
current protection of some of the 10 known populations in the S and one in the N (unprotected),
past, present and projected future decline combine to have Neocallitropsis pancheri listed as
Endangered (EN) on the IUCN Red List, up one category from the 1998 assessment.

Taxaceae

The presence of the family Taxaceae in New Caledonia seems to be an anomaly. The Taxaceae
has an almost exclusively Northern Hemisphere distribution (see map GTC-15 on p. 20) except for
one representative, Austrotaxus spicata on Grande Terre, New Caledonia. Austrotaxus is a mono-
specific genus and there is no doubt concerning its taxonomic position, confirmed both by mor-
phology and DNA. The question to be asked and answered is: how did it get to be here? (­Farjon,
2008, pp. 157–158). Seeds of the members of Taxaceae are dispersed by animals, usually birds or
less commonly mammals. Dispersal is one possibility. The fossil record of Taxaceae goes back to
the Early Jurassic of Europe and certain records of the family based on fertile material are all from
the Northern Hemisphere. However, fossil wood from the Eocene of Argentina recently described
as a new species in Taxaceoxylon may belong here or with Cephalotaxaceae (Brea et al., 2009).
Phylogenetic analysis placed Austrotaxus spicata as a sister clade to Pseudotaxus chienii + Taxus
using plastid and nuclear DNA (S. Mathews, unpublished data). Divergence time estimated for
the split appears to have been sometime in the Late Cretaceous. This evidence indicates a time
of origin when New Caledonia, as part of the Australian part of Gondwana, was separated from
 australasia 405

Asia by the Tethys Ocean, making a dispersal event very unlikely. The possibilty that ­Taxaceae
were once present in the Southern Hemisphere, and that Austrotaxus originated there, seems
more plausible.

Map AU-94. Austrotaxus spicata [n=28,

m=26, h=6, 1914–2003]

Austrotaxus spicata, a distant member

of the family Taxaceae with just one
species, occurs in several mountain mas-
sifs on Grande Terre, most abundantly
in the Table Unio ‒ Plateau de Dogny
area of Province Sud and in the Massif
du Panié in Province Nord. Another
well known locality is Mt. Canala on the
E coast. This species occurs in rainforest
at altitudes between 500 and 1350 m on
ultramafic rocks (serpentine) as well as
acidic micaschist (Mt. Canala, Massif du
Panié). It is somewhat enigmatic that it
has not been collected from the southern part of the main island, where similar habitat occupied
by similar associated trees and shrubs occur. When sterile, it may resemble Podocarpus spp., but
this handicap for detection similarly applies to the northern 2/3 of the island.

Podocarpaceae

The family Podocarpaceae is well represented in New Caledonia, with eight genera and 19 spe-
cies. Only one genus, Parasitaxus, is endemic, while Acmopyle occurs here and in Fiji, further E
in the SW Pacific (map AU-119). The other genera are widespread, i.e. Dacrydium, Dacrycarpus
and Podocarpus, or they have a highly disjunct distribution, i.e. Retrophyllum (SW Pacific, Male-
sia, South America), Prumnopitys (SW Pacific, Australia, Central America, South America) and
Falcatifolium (New Caledonia, Malesia). The link through two genera to South America is particu-
larly interesting because we have fossil evidence for Prumnopitys and Retrophyllum in Australia
(including Tasmania) and New Zealand. Some fossils of Prumnopitys found in New Zealand date
back to the Palaeocene (ca. 60 Ma) while others are Miocene (ca. 20 Ma) in age, and the genus
still occurs there. These combined facts appear to be convincing evidence against the hypothesis
of a ‘total drowning’ of New Caledonia during the Palaeocene (ca. 60 Ma) and New Zealand dur-
ing the Oligocene (ca. 30 Ma) as well as contradicting the idea that all terrestrial biota (or their
ancestors) arrived by dispersal after the drowning (e.g. Pole, 1994). Such hypothetical dispersal
origins require it to have happened at least three times for Prumnopitys (also from South America,
or perhaps to South America?). When one such cross-ocean migration is an unlikely but not
impossible chance event, a second or third arrival by the same taxon must be ruled out as too
improbable. We concur with the view that conifers demonstrate that neither New Caledonia nor
New Zealand were completely drowned at any time in their geological history (e.g. Knapp et al.,
2007) and that many of these conifers are indeed of Gondwanan origin.
406 australasia

Parasitaxus

Map AU-95. Parasitaxus usta [n=25, m=24, h=6, 1861–2003]
Parasitaxus usta, a parasitic conifer
that grows on the roots of its host, Fal-
catifolium taxoides, in the deep shade
of the forest in New Caledonia. Photo
credit William Baker.

Parasitaxus usta is most peculiar in that it is the only known parasitic gymnosperm. Its host is
another member of the Podocarpaceae, Falcatifolium taxoides, a small tree. Its habitat is therefore
that of its host: moist tropical montane angiosperm forest with scattered conifers, commonly
with emergent Araucaria spp. or less often Agathis spp. Parasitaxus usta grows usually in the
deep shade of a multi-layered canopy, never in full sunlight. The distribution of Parasitaxus usta
is rather scattered and, not surprisingly, largerly coincides with that of its host (map AU-96 on
p. 407). Obviously, where Parasitaxus usta occurs, Falcatifolium taxoides is also present, but not
the other way around. Its phylogenetic relationship is found to be with Lagarostrobos of Tasma-
nia and Manoao of New Zealand (Biffin et al. in Turner & Cernusak, eds., 2011). The estimated
divergence time of this clade (i.e. the separation of Parasitaxus from the other two genera) is
estimated to have taken place sometime during the Cretaceous. This is another instance of evi-
dence from conifers against dispersal to the islands after total drowning episodes during parts of
the Tertiary as hypothesized by Pole (1994) and others. Parasitaxus usta is classified as Vulnerable
(VU) on the IUCN Red List.
 australasia 407

Falcatifolium

Map AU-96. Falcatifolium taxoides

[n=45, m=45, h=3, 1861–2008]

Falcatifolium taxoides is the only

species of this genus that occurs
in New Caledonia. Five other spe-
cies occur in Malesia, and a map
(map MA-50 on p. 320) and dis-
cussion of the biogeography of
the genus is given in that chap-
ter. The New Caledonian species
is distributed across the length of
Grande Terre, but in the north-
ern part of the island it follows
the high and wet E coast and is
absent on the western side. Its
habitat is rainforest, in which it is
a small understorey shrub or tree
to 15 m tall, and it is most commonly found on mountain ridges and summits. It is indifferent to
the various types of substrate on these mountains, occurring on ultramafic rock as well as acidic
igneous or metamorphic rock.

Prumnopitys

Map AU-97. Prumnopitys ferrugi-

noides [n=28, m=28, h=7, 1876–2006]

The genus Prumnopitys has nine

species, only one of which occurs
in New Caledonia. Most species
are in Central and South America,
and the genus is mapped (map
SA-5 on p. 432) and discussed in
the chapter South America. Prum-
nopitys ferruginoides is a tree to
20 m tall of montane rainforest
occurring mainly on ultramafic
soils derived from serpentine; in
the extreme north of the island
(Mont Panié) it occurs on mic-
aschist. It is distributed across
Grande Terre, but rare in the cen-
tral part of the island; its altitude ranges from 150 to 1400 m. It is also absent from most of the
mountains along the W coast, presumably these do not receive enough rainfall, or more likely,
suffer droughts from time to time, particularly in El Niño years.
408 australasia

Acmopyle

The genus Acmopyle has two species, one in New Caledonia and one in Fiji (Viti Levu). In the
fossil record, Acmopyle is known from Australia (mainland), Tasmania, Antarctica and South
America. Could dispersal be responsible for its present distribution? The ‘pseudo-fruits’ subtend-
ing the seeds are without doubt eaten by birds, as they are colourful and succulent, but not very
tasty. The seeds are well protected by a strong and hard seed coat inside a softer epimatium,
which can also be digested, but more slowly. We do not know what species of birds eat them and
how far they carry them, but presumably at least from one mountain to another. Phylogenetic
analysis of Podocarpaceae has placed Acmopyle relatively basal in the cladogram (Biffin et al. in
Turner & Cernusak, eds. 2011) as sister to the Dacrydioid + Podocarpoid clades. The divergence
time was estimated to have been sometime during the Late Cretaceous, which is of course not at
all accurate, but earlier than 65 Ma. The oldest known fossils are from the Late Palaeocene, ca.
56 Ma and the genus became extinct in Australia in the Late Oligocene, ca. 25 Ma. This fits well
with a Gondwanan origin and a break-up of the supercontinent in the (Late) Cretaceous, with Aus-
tralia separating from Antarctica in the Palaeocene and starting to drift northward in the Eocene.
Dispersal was theoretically possible from New Caledonia to Fiji; the estimated divergence time
between the two species is from Late Cretaceous through Paleogene. The Fiji archipelago arose
as a consequence of subduction of the Pacific tectonic plate westward under the Indo-Australian
plate, creating an arc of islands ranging from Tonga and Fiji through Vanuatu to the Solomon
Islands, beginning in the Eocene ca. 40 Ma. (Colley in Gillespie & Claque, eds. 2009). The ancestor
of the Fijian species could have arrived there in the Eocene to Oligocene from New Caledonia.

Map AU-98. Acmopyle pancheri [n=48, The seed cone of Acmopyle pancheri consists of an edible,
m=47, h=5, 1869–2005]
red and succulent receptacle and a hard, inedible seed cov-
ered in a leathery epimatium. Birds swallow the whole and
may carry the seed a fair distance before it drops out. Dis-
persal by birds is probably responsible for the presence of
the genus, with a distinct species, in Fiji, having travelled
from New Caledonia millions of years ago. Other evidence
points at vicariance as the best explanation for the pres-
ence in New Caledonia. Photo credit Bedgebury Pinetum
(Forestry Commission).

Acmopyle pancheri is distributed across the length of Grande Terre, from the Plaine des Lacs in
the south to Mont Mandjélia in the north of the island. In the far south, it comes down as low
as 50 m a.s.l., but in most of Grande Terre it is a montane species, often on ridges and summits,
mostly between 700 and 1200 m. It is a small to medium size tree in rainforest, or in the southern
part of the island in the ecotone between forest and ‘maquis minier’ on lateritic soils. It appears
to be indifferent to rock type, as it occurs on ultramafic rocks as well as acidic micaschist. High
rainfall requirements keep it away from the W coast, most notably in Province Nord.
 australasia 409

Dacrycarpus

Map AU-99. Dacrycarpus vieillardii

[n=36, m=34, h=5, 1861–2005]

The genus Dacrycarpus has nine

species and is widely distributed
from SW China to New Zealand,
but it is absent in Australia. This
genus is mapped (map MA-43 on
p. 316) and discussed in the chap-
ter Malesia. Only one species is
present in New Caledonia: D. vie-
illardii. This species occurs mostly
along streams across Grande
Terre from the Plaine des Lacs in
the south to the northern part,
from near sea level to ca. 900 m. It
is more or less limited to regions
with ultramafic rocks, hence its
virtual absence in the far north of the island. However, it seems also absent from several of the
ultramafic massifs there. Its relative scarcity in the north may not be entirely genuine but partly
an artifact of collecting activity. Other lacunae are the drainages from Mt. Humboldt, Mt. Mou,
Mt. Do, Table Unio and other mountains with ultramafic formations in the central part of Prov-
ince Sud. These locations are well collected, so there are probably unknown ecological reasons for
the absence of Dacrycarpus vieillardii there; see also Podocarpus lucienii (map AU-104 on p. 412).

Retrophyllum

The genus Retrophyllum has five species, two in New Caledonia, one in Malesia, the Santa Cruz
Islands and Fiji, and two in South America. The distribution of this genus therefore consists of two
very disjunct regions, E Malesia to tropical SW Pacific, and tropical Andes in South ­America. We
have seen this disjunct link between the SW Pacific and South America in several other ­genera,
e.g. Araucaria (map AU-74 on p. 391) and Prumnopitys (map SA-5 on p. 432). If we include the
fossil record we may add Dacrycarpus and Acmopyle, still extant in the SW Pacific but extinct
in South America (Farjon, 2008, p. 173) Many of these have been found in the fossil record of
Tasmania, the last link with Antarctica before Australia began drifting north in the Eocene (Hill &
Brodribb, 1999). It would appear that no long distance dispersal across the Pacific (the direct
route) has ever taken place; the cradle of the southern conifers lies in fossil beds under the ice
of Antarctica.
410 australasia

Malesian Pacific

Tropical Andean

Map AU-100. The distribution of Retrophyllum in two very disjunct regions. As exemplified in several other
genera the link between the SW Pacific and South America is a recurrent biogeographical phenomenon in
the southern conifers. In this case the distribution in South America is tropical-Andean, but the southern
Andes is home to several other conifer genera with this disjunction. Likewise, the distribution of Retrophyl-
lum in the east is Malesian-Pacific, but New Zealand and Tasmania can also be involved on that side of the
Pacific Ocean. Including New Guinea in Malesia, all form part of or are accreted to the drifted fragments
of the southern supercontinent Gondwana.

Map AU-101. Retrophyllum comptonii

[n=44, m=43, h=8, 1902–2008]

Retrophyllum comptonii is a tree

to 30 m tall occurring in montane
rainforest throughout Grande
Terre, at altitudes of 600–1450 m.
In the far south it has been found
near sea level at Port Boisé, close
to a small population of R. minus.
Retrophyllum comptonii occurs
on serpentine and other ultra-
mafic rocks as well as on acidic
micaschist (e.g. Mont Panié).
This species is shade tolerant
and can linger under canopy,
growing slowly, until an oppor-
tunity arises to fill a small gap
and grow taller. In this way, several podocarpaceous conifers can compete with angiosperms in
tropical forests (Enright & Jaffré in Turner & Cernusak, 2011). Common associated conifers are
­Araucaria spp. and Agathis spp. (Araucariaceae), Acmopyle pancheri, Falcatifolium taxoides, and
to a lesser extent Podocarpus spp. (Podocarpaceae). Conifers predominate on ridges, angiosperms
on slopes, but R. comptonii is successful even as a solitary tree growing up among angiosperms.
The ‘tortoise and the hare’ ecological strategy (Bond, 1989) does not apply to all conifers. Per-
haps the peculiar phyllotaxis of the amphistomatic leaves enables them to compete successfully
with the ‘more efficient’ leaves of the faster growing angiosperms.
 australasia 411

Map AU-102. Retrophyllum minus Retrophyllum minus in the Rivière des Lacs, Plaine des Lacs, New
[n=23, m=23, h=9, 1861–2000] Caledonia. These dwarfed trees germinate in mud and grow in
running water; they are the only known true rheophytes among
conifers. Photo credit C. N. Page.

Retrophyllum minus is a small tree or shrub to 3 m tall, growing in or immediately alongside

streams and lakes in the far S of Grande Terre, mainly in the Plaine des Lacs and the middle
drainage of the Yaté River, and in a few localities near the S coast of the island. It is a true
rheophyte (the lake occurrences are on small lakes that are part of the Rivière des Lacs drain-
age). Rheophytes are plants that grow in running water. The buoyant seeds are easily transported
by water and germinate in mud temporarily falling dry with fluctuating water tables in lakes
and streams. This dwarfed tree is therefore limited to water courses. It is also limited to the
lowlands and occurs from near sea level to 240 m. The streams drain a large area of ultramafic
rock, mainly laterites, and some hills and low mountains of weathered serpentine. The streams,
particularly after heavy rains, can be contaminated by partly dissolved metals; one creek where
Retrophyllum minus occurs is known as Pernod Creek. Like the French alcoholic beverage, which
turns milky when diluted with water, this stream aquires that colour after rain. This species is
listed as Endangered (EN) on the IUCN Red List, because of its specialized habitat which can be
negatively impacted by nickel mining in various ways, including drainage of the Plaine des Lacs
should mining ever start there on a large scale. Only a tiny part of the entire population is in a
protected area.

Podocarpus

Seven species of Podocarpus are recognized in New Caledonia (Farjon, 2010a); they range from the
lowlands to the mountains and from ‘maquis minier’ and riparian habitats to rainforest. A few are
widespread, some others have limited distributions. Some are shrubs, most are trees of moder-
ate size. Only two species from New Caledonia were included in a recent phylogenetic analysis
(­Biffin et al. in Turner & Cernusak, eds. 2011) using molecular (DNA) data, P. sylvestris and P. gnid-
ioides. They appeared in two different clades that divide the genus, with P. sylvestris belonging
to subgenus Foliolatus and P. gnidioides in subgenus Podocarpus. As the entire ­pantropical genus
­apparently divides into these two subgenera, it is clear that the seven species of Podocarpus in
New Caledonia derive from two ancestors that must have arrived after this major split in the
genus occurred. This split is estimated to have occurred sometime in the Paleogene, i.e. after
65 Ma, when New Caledonia had already separated from Australia. If this timing is correct, a
minimum of two dispersal events is involved in the origin of the species presently occurring in
New Caledonia. It would be interesting if the phylogenetic relationships among all seven of these
species could be investigated.
412 australasia

Map AU-103. Podocarpus sylvestris [n=47,

m=45, h=5, 1861–2008]

Podocarpus sylvestris is widespread and

is distributed acorss the length of
Grande Terre as well as on the Île des
Pins. It is a small to medium size tree of
montane rainforest, sometimes appear-
ing in the ecotone to ‘maquis minier’
where it remains a small tree like the
other species on ultramafic soil. In
regions dominated by maquis due to
soil and fires, it retreats with the forest
into sheltered ravines or gullies on rain-
exposed SE or E slopes. It is not restricted
to ultramafic substrates and also occurs
on acidic micaschist in the northern part of the main island. Its altitude range is 150 to 1200 m. In
dense rainforest it can reach 30 m, but most trees remain smaller and in the lower canopy, where
they can tolerate shade and come to reproduction. Podocarpus sylvestris is therefore a tree with
a scattered but common appearance in the forests of New Caledonia.

Map AU-104. Podocarpus lucienii [n=31,

m=31, h=9, 1942–2005]

Podocarpus lucienii is distributed

unevenly on Grande Terre, with com-
mon occurence in what is known geo-
logically as the ‘Massif du Sud’, i.e. the
Plaine des Lacs and surrounding hills
and mountains, and a more scattered
distribution in the northern half of the
island. Some other species in the family
Podocarpaceae have a similar distribu-
tion pattern, e.g. Parasitaxus usta (map
AU-95 on p. 406), Prumnopitys fer-
ruginoides (map AU-97 on p. 407) and
Dacrycarpus vieillardii (map AU-99 on
p. 409). They are absent or rare in the south-central part of the island. Habitats suitable for these
species seem to be widespread there and an explanation for this distribution pattern is wanting.
Podocarpus lucienii occurs on both ultramafic and acidic rock as a small understorey tree in rain-
forest at 250 to 1350 m a.s.l. It is often more common on ridges than on densely forested slopes.
This species is listed as Endangered (EN) on the IUCN Red List due to deforestation and threats
from nickel mining.
 australasia 413

Map AU-105. Podocarpus decumbens [n=8, m=8,

h=5, 1949–1967]

Podocarpus decumbens has a very limited dis-

tribution. Most of the few herbarium collec-
tions known were made on the Montagne des
Sources; two nearby localities are Pic du Rocher
and an unnamed ridge above the Rivière Bleue.
These collections date from 1949 to 1967 and no
recent collections have been added to the main
institutional herbaria. This species is a pros-
trate, decumbent shrub, spreading over rocks,
but sometimes semi-erect when supported by
low shrubs. Some have claimed that it is a
climber, but there are no apparent structures in
the stems such as can cause twining, or other
morphological aids to climb, and we can safely say that conifers have not produced plants with
a true climbing habit. It is capable of layering, i.e. forming secondary roots on decumbent branches
or stems. Although it occurs on remote ridges at 800–1000 m and mostly within protected areas,
the fact that it has not been collected for nearly 50 years and that its populations are very small
and vulnerable to fires, which have increased due to carelessness from hikers, has caused this
species to be listed as Critically Endangered (CR) on the IUCN Red List.

Map AU-106. Podocarpus polyspermus [n=8,

m=8, h=5, 1962–1977]

Podocarpus polyspermus is also rare and

few herbarium collections have been
made since it was named and described
in the Flora of New Caledonia (De
Laubenfels, 1972). It occurs in the central
part of Grande Terre on several moun-
tains at altitudes between 650 and 950 m
in low forest or in transitional situations
(ecotone) to ‘maquis minier’ on ultrama-
fic substrates. There it grows to a small
tree 5–15 m tall, joining the canopy or
staying somewhat below it in taller for-
est habitat. Although still imperfectly
known, its range is undoubtedly limited.
In several places it occurs on mountains
where nickel mining is destroying its
habitat; this species is listed as Endan-
gered (EN) on the IUCN Red List.
414 australasia

Map AU-107. Podocarpus longifoliolatus

[n=17, m=17, h=4, 1870–1982]

Podocarpus longifoliolatus is uncommon,

collected from only five mountains on
Grande Terre; most specimens in her-
baria came from Mt. Mou (counting
from the S on Grande Terre the second
dot on the map). This mountain is easy
to reach from the main road along the
E coast of the island. It is also present on
Pic N’Ga on the Île des Pins, at a much
lower altitude (150–180 m) than the
other localities, where it occurs above
1000 m. This species has only been found
on ultramafic rock, mostly serpentine or
its eroded derivatives. As with other species, its size is dependent on the vegetation surrounding
it, but trees to 20 m tall have been encountered. Podocarpus longifoliolatus is listed as Endangered
(EN) on the IUCN Red List.

Map AU-108. Podocarpus novae-caledoniae Podocarpus novae-caledoniae is a shrub or rarely a small

[n=37, m=30, h=3, 1861–2005] tree growing exclusively on ultramafic rocks in the south-
ernmost part of Grande Terre, New Caledonia, as well as
on the Île des Pins. Photo credit Aljos Farjon.

Podocarpus novae-caledoniae is nearly limited to the southern part of Grande Terre known by
geologists as the’ Massif du Sud’, where it is common along rivers and creeks in the lowlands and
on lower mountains to 700 m. It is not strictly riparian and can be found on slopes away from
streams. The vegetation bordering streams in the Plaine des Lacs is generally lusher than that on
the parched terrain between streams, it may also be somewhat less prone to burn. The recently
described species P. beecherae was said to be non-riparian but is here considered a taxonomic
synonym (Farjon, 2010a). Podocarpus novae-caledoniae is a shrub occurring only on ultramafic
substrates, in ‘maquis minier’ vegetation. Among trees such as Agathis ovata it can occasionally
become a small tree 4–6 m tall. A disjunct occurrence is on the Île des Pins, where it is found
on Pic N’Ga together with P. longifoliolatus and P. sylvestris. This small mountain is an isolated
outcrop of serpentine and thus ultramafic, hence the occurence of these species there.
 australasia 415

Map AU-109. Podocarpus gnidioides [n=24, m=22,

h=6, 1862–2005]

Podocarpus gnidioides is distributed in the

southern part of Grande Terre, from the Forêt
de Sailles in the north to Mont Dore in the
south. It is a small-leaved shrub of the higher
mountains above 600 m and has been found
on the summit of Mt. Humboldt at 1618 m. It
occupies slopes and ridges with low vegetation
on serpentine or other ultramafic rocks, in sun-
exposed situations but with very high rainfall.
It is similar in habit and morphology to some
subalpine species of Podocarpus in Australia
and New Zealand and may be related to these.
More comprehensive taxon sampling in future phylogenetic analysis is needed to confirm this;
such research should include all species of Australasia and several of Malesia. Its limited range is
somewhat mysterious because mountains with sufficient height and ultramafic rocks do extend
further NW than where this species has been collected. Its dispersal mechanism is similar to other
species, but we only know the generalities of this and not the specific limitations that perhaps
apply.

Dacrydium

The genus Dacrydium has five species in New Caledonia; one is a recently described hybrid
(nothospecies). Dacrydium is widespread from mainland Southeast Asia through Malesia and the
Southwest Pacific to New Zealand. Its greatest diversity is in Malesia and a map (map MA-31 on
p. 309) of the genus with discussion of its biogeography is presented in that chapter.

Map AU-110. Dacrydium balansae

[n=40, m=39, h=6, 1864–2005]

Dacrydium balansae is a small

tree occurring mostly on ultrama-
fic soils derived from serpentine
at elevations from near sea level
to 800–900 m. It is distributed
across Grande Terre and follows
the ultramafic mountain massifs.
It is therefore not found in the
NE of the island, where acidic
metamorphic rocks prevail. Other
areas where it is absent are in
the central part of the island,
where large areas are made up
of Upper Palaeozoic to Mesozoic
sediments. Absence in the far SE
(Plaine des Lacs and surrounding hills and mountains) is not related to substrate, as this is also an
ultramafic area and the species is found at lower altitudes. It is replaced there by D. araucarioides
(map AU-112 on p. 416).
416 australasia

Map AU-111. Dacrydium lycopodioides [n=18, m=18,

h=3, 1864–2005]

Dacrydium lycopodioides is a relatively rare

species, but its distribution has been found
to be more extensive than was indicated by
a map presented in the Flora of New Caledo-
nia (De Laubenfels, 1972), in which most of
the herbarium collections cited came from
Mt. Mou. Dacrydium lycopodioides occurs in
higher mountains at altitudes between 800 m
and 1400 m and in some wooded ravines in the
interior at slightly lower elevation. It extends
from Mt. Nakada in the NW to Mt. Mou and
the Haute Ouinné River in the SE; all loca-
tions are in Province Sud. In sheltered for-
est patches it may attain 25 m but usually it
remains a much lower tree on mountain ridges. This species is also limited to serpentine and its
derivatives, i.e. ultramafic rocks, in common with other species of Dacrydium in New Caledonia.
Narrow-leaved conifers like Dacrydium are less tolerant of shade and therefore tend to ‘retreat’
to the less favourable, ultramafic sites or to ridges, where large-leaved angiosperms are usually
absent as there are not sufficient nutrients available to sustain these.

Map AU-112. Dacrydium araucarioides [n=35,

m=34, h=5, 1861–2005]

Dacrydium araucarioides in the Forêt du Mois de Mai,

Province Sud, New Caledonia. This species is common
in ‘maquis minier’, a species-rich, open shrub vegeta-
tion on highly metalliferous, ultramafic substrates that
covers large areas, particularly in the southern parts of
Grande Terre. This vegetation is subject to fires and
the soil is deficient in basic minerals needed for plant
growth. This conifer is well adapted to these adverse
conditions. Photo credit Aljos Farjon.
 australasia 417

Dacrydium araucarioides is limited to the southern half of Grande Terre and most common in the
‘Massif du Sud’ at the southern end of the island. It reaches its northernmost station near Poro on
the E coast and is not found on the W coast, where it is too dry. Dacrydium araucarioides occurs
from near sea level to just under 1000 m and at high elevations it is found on steep ridges and
rocky summits, where competition from large trees is limited to a few trees, usually a species of
Araucaria. It occurs on ultramafic substrates, especially on laterite soils and ‘ironstone’ (‘cuirasse
de fer’) on the Plaine des Lacs and in the surrounding hills. This species occurs as solitary small
trees or sometimes in gregarious stands, the tallest individuals rising above the lower vegetation
(‘maquis minier’). In these harsh environments, prone to fire and with an almost impenetrable
duricrust soil, this little tree is a very slow grower and develops a characteristic candelabra-shaped
crown. As with other podocarps, the small seeds of this species are dispersed by birds. Perhaps it
recolonises from surviving trees in the area after a fire, but no specific studies have been under-
taken to investigate how this species manages not only to survive but to remain common in an
inhospitable environment.

Dacrydium guillauminii on the water edge of the Rivière des

Map AU-113. Dacrydium guillauminii [n=11, Lacs, Province Sud, New Caledonia. This small conifer is
m=11, h=5, 1948–1996] resistant to toxic water containing heavy metals and occu-
pies a narrow strip between high and low water lines. Photo
credit Alexander Schmidt.

Dacrydium guillauminii has a very limited distribution, due to a peculiar habitat which it more or
less shares with Retrophyllum minus (map AU-102 on p. 411). It is found on the banks of a few small
rivers and creeks and on two lake shores in the Plaine des Lacs, in the far SE of Grande Terre.
The main river is the Rivière des Lacs with a few tributaries, originating in the Grand Lac and
the Lac en Huite. Dacrydium guillauminii is a riparian species, ocurring in the zone between high
and low water lines subject to both flooding and drought. Its seeds germinate in moist shores,
but not in water. The Rivière des Lacs drains ultramafic hills and plains and when in flood can be
charged with highly metalliferous silt that is poisonous to all but the adapted species of plants.
A high level of tolerance allows these two conifers to occupy this narrow niche to the exclusion
of most other plants. Just metres away on higher ground, the ‘maquis minier’ vegetation begins
and there other conifers and angiosperms prevail. Dacrydium guillauminii is very rare and its
sub-populations are small and vulnerable to disturbance, including fire, alteration of river flow
through damming, and potential mining in the area. Only one small nature reserve at the Chutes
de la Madeleine protects this species, which is listed as Critically Endangered (CR) on the IUCN
Red List.
418 australasia

Map AU-114. Dacrydium ×suprinii [n=3, m=3, h=3,

2005]

For several years, observers of the remarkable

conifer dwarf forest in the drainage of the Riv-
ière des Lacs have observed a few individuals
that seemed to combine morphological charac-
ters from two species, Dacrydium araucarioides
and D. guillauminii. It appears to grow with the
putative maternal parent, D. guillauminii. Sur-
veys conducted more recently have found indi-
viduals along the Rivière des Lacs and on the
shore of Grand Lac where this river originates.
A detailed morphological study (Knopf et al.,
2007) confirmed the hybrid origin as a fixed
nothospecies, but further work on the genetics
of all three taxa is needed to establish its status
beyond doubt as more than just an F1 hybrid.
Very few herbarium collections of Dacrydium ×suprinii have been made to date and only three
localities can therefore be shown on the map. This taxon is not listed on the IUCN Red List pend-
ing its definitive taxonomic status; if it were considered a species it would be Critically Endan-
gered (CR).
 australasia 419

The ‘interarc’ islands of the Southwest Pacific

Geography

We consider in this section the remainder of islands in the SW Pacific Ocean that are situated
on the Indo-Australian tectonic plate, New Caledonia and New Zealand having been discussed
elsewhere in this chapter. These islands form the following archipelagoes: Fiji, Solomon Islands,
Tonga, Vanuatu and the Wallis & Futuna Islands. Within these archipelagoes, smaller groups of
islands relevant to conifers occur, such as the Santa Cruz Islands. Bougainville, although part of
the Solomon Islands Archipelago, is an autonomous province of Papua New Guinea and is treated
under Malesia. Norfolk Island and Lord Howe are isolated islands in the Tasman Sea situated
between New Caledonia, Australia and New Zealand. The region is geologically characterised
by convergence between the Pacific and Indo-Australian plates. The archipelagoes are situated
on the Indo-Australian plate side of a complex arc of subduction zones, where the Pacific plate
slowly disappears under the continental plate. Generally, this subduction zone forms a huge arc
from New Zealand NE to Tonga and Samoa and from there WNW to Vanuatu. Inside this arc
there are smaller subduction zones, some no longer active. The archipelagoes were formed as
a result of tectonic movements of the two plates, some as a marginal side effect of subduction
(Tonga), while others arose on large platforms (Fiji), but all are part of a continuous volcanic
arc associated with the convergence of these two plates of the Earth’s crust. These islands are all
‘oceanic’ in the sense that they arose from the ocean floor during the Tertiary and were never, as
far as we know, connected to a continent. Norfolk Island and Lord Howe differ in that they are
situated on the Norfolk Ridge and the Lord Howe Rise, sections of a now submerged continental
platform that was once part of a landmass (‘Zealandia’) conneting New Zealand and possibly New
Caledonia with the eastern part of Gondwana. However, Norfolk Island is of more recent volcanic
origin, beginning ca. 3 Ma. Most of the islands are small and the largest are Viti Levu (Fiji) with
10,389 km², Vanua Levu (Fiji) with 5,587 km², Guadalcanal (Solomon Islands) with 5,302 km² and
Espiritu Santo (Vanuatu) with 3,955 km². Several are mountainous, with the highest points Mt.
Tomanivi on Viti Levu at 1324 m, Mt. Tabwemasana on Espiritu Santo at 1,879 m and Mt. Popo-
manaseu on Guadalcanal at 2,449 m. The climate on all islands is tropical, with SE trade winds
from April to October and cyclones occurring from November to March under influence of the
NW monsoon.

Conifers on the ‘interarc’ Islands

Families Genera Species Species + infra Countries with Area of occupancy Endemic taxa
conifers in km²
2/8 7/70 19/615 20/794 5/6 918/62,180 11
Numbers after ‘/’ are global or continent totals.

The ‘interarc’ islands in the SW Pacific here considered have a modest diversity of conifers, with
two families, seven genera and 19 species + one variety. The Wallis & Futuna Islands and Lord
Howe have no native conifers. Endemism is moderate at 55% of taxa for the region as a whole
(nine species also occur outside the region in Malesia), but is higher when the archipelagoes are
considered separately (see map AU-3 on p. 342). The two families are Araucariaceae and Podo-
carpaceae. In Araucariaceae, two genera are present, Agathis with two species and the single spe-
cies Araucaria heterophylla endemic on Norfolk Island. Podocarpaceae is the larger group in the
region, with five genera, 16 species and one variety. Of these genera, Podocarpus is represented
with eight species + one variety, Dacrydium with five species, and Acmopyle, Dacrycarpus and
Retrophyllum with one species each. Given the origin of the islands, the presence of these conifers
must go back to dispersal events, involving the species currently found there, or their ancestors.
420 australasia

Island-hopping along the arc from the Malesian region via New Guinea seems the most likely
route for several species or their ancestors (Doyle, 1998). Nine species, Dacrycarpus imbricatus,
Dacrydium beccari, D. magnum, D. nidulum, D. xanthandrum, Podocarpus glaucus, P. neriifolius,
P. pilgeri and Retrophyllum vitiense are more widely distributed in Malesia (three of these are
also native in mainland SE Asia) and reach the Solomon Islands (Dacrydium beccari, D. magnum,
D. xanthandrum, Podocarpus glaucus, P. pilgeri) and Vanuatu and/or Fiji (Dacrycarpus imbrica-
tus, Dacrydium nidulum, Podocarpus neriifolius and Retrophyllum vitiense). They are mapped and
discussed in the chapter Malesia. Podocarpus insularis is not strictly endemic to the ‘interarc’
islands here defined because it extends to the nearby Louisiade Archipelago, the D’Entrecasteaux
Islands, the E part of New Guinea and New Britain. Some of the species of the ‘interarc’ islands
are more closely related to species in New Caledonia, e.g. Acmopyle sahniana and Dacrydium
nidulum (Biffin et al. in Turner & Cernusak, eds. 2011). It is very interesting to note that none
of the widespread species occur in New Caledonia, which gives further proof of its ‘deep time’
isolation and the separate evolution of its conifers. Without further and more comprehensive
phylogenetic analyses of the Podocarpaceae in particular, the routes and direction of migration
remain mostly speculative. While the species in Podocarpaceae are dispersed by birds and can
presumably be taken as seeds across the sea, the mode of dispersal across the same for the three
species in Araucariaceae remains enigmatic. Seeds of Araucaria are fused with the cone scales
and fall to the ground with them; the animals that eat them are not known to swim in the sea for
long. Seeds of Agathis are winged but do not appear to travel great distances, however they may
be transported by water (Doyle in Farjon & Page, comp. 1999).

Tonga

Map AU-115. The distribution of conifers on the ‘interarc’ islands of the SW Pacific. In this chapter these
are the archipelagoes of the Solomon Islands, Vanuatu, Fiji and Tonga and Norfolk Island (the latter not
shown on the map).
 australasia 421

Araucariaceae

Two species of Agathis and one species of Araucaria occur on the islands here discussed. One spe-
cies of Agathis (A. macrophylla) is widespread in the region and occurs in three archipelagoes. The
other species, Agathis silbae and Araucaria heterophylla are known from a single island each.

Map AU-116. Agathis macrophylla

[n=47, m=37, h=5, 1852–1999]

Agathis macrophylla occurs natu-

rally in Fiji on Kadavu Island, Viti
Levu and Vanua Levu; in the Sol-
omon Islands in the Santa Cruz
Group on Utupua Island and Van-
ikoro Island; and in Vanuatu on
Anatom [Aneityum], Erromango
and Tanna. This species has been
planted elsewhere in the archipel-
agoes of the SW Pacific, but such
plantations are not mapped. The
erratic natural distribution of this
species is apparent from the map. It is particularly striking that it is absent from all the islands
of Vanuatu between Erromango in the south and Vanikoro in the Santa Cruz Group in the north.
There is abundant suitable habitat on the intervening islands. This habitat is tropical evergreen
broadleaf rainforest from the lowlands to ca. 900 m a.s.l. Logging has negatively affected this spe-
cies especially in the Santa Cruz Group, where on Utupua Island trees in the wild may now have
effectively disappeared (Doyle in Farjon & Page, comp. 1999). Exploitation in Fiji and Vanuatu has
also caused decline, but in no cases has this led to the disappearance of the species from islands
in these two archipelagoes. Could it have become extinct on the islands of Vanuatu where it is
not found? It is apparently capable of competing with other trees in the forest, where large trees
of Agathis are usually emergents (Whitmore, 1966). It grows in soils derived from volcanic rocks,
but all the main islands are of volcanic origin. Could climatic fluctuations in the geological past
have had an effect? Or is it a case of erratic chance dispersal? Agathis is unlikely to be a long
distance disperser; although its seeds have a wing and detach from the scale when the seed cone
disintegrates, short distance island-hopping would be a more plausible event. Past extinction is
therefore more likely to have occurred, but the causes of this remain unknown. Agathis macro-
phylla is listed as Endangered (EN) on the IUCN Red List.
422 australasia
Map AU-118. Araucaria heterophylla
[n=14, m=11, h=5, 1788–1968]
Map AU-117. Agathis silbae [n=10, m=9, h=6,
1979–1987]
Agathis silbae is a relatively unknown but dis-
tinct species (the distinction, as with most spe-
cies of this difficult genus, is in the morphology
of the pollen cones) occurring on Espiritu Santo,
the largest island in Vanuatu. The first trees
described were planted in villages along the
W coast and only later, in 1986 was it found in
the mountain forests of the interior. The her-
barium specimens taken from planted trees
have not been mapped. In the tropical ­forest
on the W side of the island it occurs as an
emergent; sometimes very large trees to 4 m in
diameter and 40 m tall have been reported. Its
ecology is not well known but may be similar to
that of A. macrophylla in Fiji.
Araucaria heterophylla on Norfolk Island in semi-natural habitat
(with grasses mostly introduced and broadleaf trees disappeared).
Larger trees may be remnants of the original vegetation, which
was subtropical broadleaf rainforest with the Norfolk Island pines
as emergents, most abundant near the coast of the island. Photo
credit © thinboyfatter, downloaded from Wikimedia Commons.
 australasia 423

Araucaria heterophylla, the Norfolk Island pine is known the world over as an ornamental tree,
either grown in a pot indoors or planted in warmer climes in gardens, parks and along roads, usu-
ally near the sea coast. Yet it is native only on Norfolk Island, a tiny spot of only 34.6 km² in the
Pacific Ocean halfway between New Caledonia and New Zealand and ca. 1500 km E of Australia.
When discovered by James Cook in October 1774 much of the island was covered in subtropical
rainforest, in which A. heterophylla was an emergent tree, similar to situations still present in
New Caledonia, 700 km to the north. Only scattered stands of the araucarias are left while the
rainforest, once rich in tree ferns, has mostly disappeared. Planting has also been frequent in
more recent times, and it is not easy to distinguish without doubt if trees are remainders of the
forest, now often surrounded by a sward of (introduced) grass, or were planted. Also, on this scale,
herbarium collections and their often less precise locality information are insufficient to produce
a map of any detail indicating where on Norfolk Island A. heterophylla may still occur as a natural
tree. The dots on the map are mostly indicative of localities. Araucaria heterophylla is listed as
Vulnerable (VU) on the IUCN Red List.

Podocarpaceae

Here we present maps only of those species that are endemic to the ‘interarc’ islands, excluding
those with a wider distribution. In Podocarpaceae, these are Acmopyle sahniana, Dacrydium nau-
soriense and five species of Podocarpus + one variety.

Map AU-119. Acmopyle sahniana [n=8, m=8, h=6,

1927–1994]

Acmopyle sahniana occurs in Fiji, where it is

only known from three locations, Mt. Evans
Forest Park (Mt. Koroyanitu) which is a pro-
tected area, Mt. Vakarongasiu [Mt. Nakorolo]
and the Korobasabasaga Range (not shown), all
on Viti Levu, the largest island. The only other
species in this genus occurs in New Caledonia
(map AU-98 on p. 408); nothing has been found
in Vanuatu, the archipelago lying between Fiji
and New Caledonia. It cannot be ruled out
that it occurs there too, because a new local-
ity on Viti Levu (Korobasabasaga Range) was
discovered in the 1990s after much searching,
and some of the islands in Vanuatu have been
incompletely surveyed botanically. Acmopyle sahniana is extremely rare, as fewer than 100 mature
trees are known to occur in the three populations so far found, with the one on Mt. Koroyanitu
now possibly extinct (Doyle in Farjon & Page, comp. 1999). It is a small tree to 12 m tall, often
looking spindly with few branches. This species is restricted to low rainforest on mountain ridges
and summits at altitudes between 375 and 950(–1050?) m in an extremely humid tropical climate.
Acmopyle sahniana is listed as Critically Endangered (CR) on the IUCN Red List.
424 australasia

Map AU-120. Dacrydium nausoriense [n=14, m=14,

h=5, 1947–1973]

Dacrydium nausoriense is known from five

localities in Fiji, three on Viti Levu and two on
Vanua Levu, based on herbarium collections. It
occurs in tropical evergreen rainforests domi-
nated by angiosperms and is fairly common in
these, at least in the Nausori Highlands on Viti
Levu. However, these forests have been reduced
to remnants in these highlands and elsewhere,
and pasture with largely introduced grasses as
well as secondary scrubland has taken their
place. The herbarium collections used for this
map do not date from later than the 1970s and
are probably incomplete. More focused collect-
ing may also result in more localities, if it could be undertaken. On the other hand, ongoing log-
ging and deforestation and the lack of adequately protected forest reserves may have caused local
extinctions. This species is listed as Endangered (EN) on the IUCN Red List.

Map AU-121. Podocarpus affinis [n=8, m=7, h=7,

1860–1973]

Podocarpus affinis has been collected in Fiji in

four localities in the South-central part of Viti
Levu and on Ovalau Island just E of Viti Levu.
Voma Peak is the best known locality, where
several herbarium collections were made. As
with the previous species, more extensive
collecting could yield new localities, but the
argument from especially deforestation also
applies, i.e. some known localities may no lon-
ger have this species. None of the collections
used to produce this map is more recent than
the 1970s. This does not mean that this species
has disappeared from any of these localities, as
the collecting frequency has been very low, but it could have gone given the ongoing deforesta-
tion. More botanical collecting is urgently needed in Fiji. Podocarpus affinis occurs as a medium
size tree in low rainforest on high ridges and mountain summits between 900 and 1200 m a.s.l.
Its status on the IUCN Red List is Vulnerable (VU).
 australasia 425

‘Eua

Map AU-122a. Podocarpus pallidus [n=9, Map AU-122b. Podocarpus pallidus in ‘Eua

m=9, h=5, 1855–1990] in Uta Vava’u

Podocarpus pallidus occurs in Tonga, on the islands of ‘Uta Vava’u and ‘Eua in two island groups
on the opposite ends of the archipelago, 325 km apart. This is on the far eastern edge of the Indo-
Australian Plate, nearly 3,300 km E of Australia and ca. 2,000 km NNE of New Zealand. Of course,
Fiji is ‘only’ 800 km away to the west, but we do not know at present to which species P. pallidus
is most closely related. By all accounts, this is an extremely isolated species of conifer. It is a
shrub or small tree, branching low and not growing taller than 10 m. Its habitat is escarpments on
the E-side of ‘Eua and low plateaux with ravines and gullies on Uta Vava’u, where it is scattered
in patches of low tropical forest, mostly composed of angiosperms. On ‘Uta Vava’u it occurs on
limestone cliffs; the altitude range is 50–250 m. No herbarium collections are known from Tonga-
tapu, Tofua or Late, other islands in the archipelago with low forested mountains and plateaux.
There may be fewer than 1000 individuals, but this is an estimate based on area of forest patches
and presumed frequency of this species. A survey on the ground is needed to establish a more
accurate estimate. On the basis of existing knowledge Podocarpus pallidus is listed as Vulnerable
(VU) on the IUCN Red List. Some part of one of the two populations is within ‘Eua National Park,
the southern half of the escarpent on the E-side of that island.

Map AU-123. Podocarpus salomonien-

sis [n=10, m=9, h=8, 1932–2008]

Podocarpus salomoniensis is the

only species of conifer endemic
to the Solomon Islands (Doyle in
Farjon & Page, comp. 1999); the
other species in Agathis, Dacry-
carpus, Dacrydium, Podocarpus
and Retrophyllum that occur there
have wider distributions, mostly
in Malesia. Podocarpus salo-
moniensis is present on three
main islands: Choiseul, Santa Isa-
bel and San Cristobal, and on
some of their ‘satellites’ such as
426 australasia

San Jorge and Pawa. Few herbarium collections were made over the years, and it seems likely
that P. salomoniensis is to be found elsewhere on these islands, as well as on other islands in the
archipelago. It occurs in rainforest and swamp forest, in the latter on ultramafic soil and accom-
panied by Casuarina and various shrubs. The altitude range is imperfectly known but could
extend from the lowlands at 10 m to 900 m or more. In the rainforest it can be a tall tree to 25 m,
but in swamp forest it is usually much smaller. This species is listed as Endangered (EN) on the
IUCN Red List.

Map AU-124. Podocarpus neriifolius var. degeneri

[n=16, m=16, h=8, 1941–1999]

Podocarpus neriifolius is the most widespread

species of its genus, occurring from Nepal and
S China to Fiji. The species is mapped in the
chapter Malesia (map MA-7 on p. 298). With
such an enormous range there will be varia-
tion in many places, but it seems that the only
consistently distinct variety occurs, together
with the nominal variety, in Fiji. It is therefore
mapped and discussed here. Podocarpus nerii-
folius var. degeneri is the most common variety
in Fiji, especially in the Nausori Highlands and
other areas in the northern half of Viti Levu.
The var. neriifolius has been found more often on the coast, but also appears to occur in the inte-
rior, at least in Viti Levu.

Map AU-125. Podocarpus insularis [n=17, m=17, h=6, 1956–1989]

Podocarpus insularis occurs from the southern peninsula of New Guinea and New Britain to Ana-
tom [Aneityum] in Vanuatu. It is present but uncommon in the Solomon Islands (including the
crater of Lake Loloru in Bougainville) and more frequent on the islands of the D’Entrecasteaux
and Louisiade archipelagoes and Woodlark Island. There is a gap in its distribution between these
islands and Erromango, Tanna (no specimen available) and Anatom [Aneityum] in Vanuatu as it
 australasia 427

has not been collected on the Santa Cruz Islands and the islands of Vanuatu between these and
Erromango. A similar but less wide gap was observed in the distribution of Agathis macrophylla
(map AU-116 on p. 421). Podocarpus insularis is a tall canopy tree of tropical evergreen rainforest,
occurring from near sea level to ca. 1900 m. It occurs on a variety of substrates, including peaty
soil and ultramafic rock; in these nutrient poor habitats it remains small or stunted and is often
accompanied by other podocarps and on ultramafics by Casuarina.

Map AU-126. Podocarpus orarius [n=13, m=13, h=3,

1929–2008]

Podocarpus orarius has been named and

described recently (Mill & Whiting, 2012) as a
result of a revision of the concept of P. spathoides
de Laub. as presented in Flora Malesiana 10 (3):
404 (De Laubenfels, 1988), where it was given a
very disparate distribution ranging from Penin-
sular Malaysia (where the type specimen came
from) via the Moluccas to the Solomon Islands.
New collecting and field observations and care-
ful comparison of types and other material led to
the conclusion that the Solomon Islands material
does not belong with P. spathoides but represents
a new species, P. orarius R. R. Mill & M. Whiting. It was also suspected that some specimens
collected on Erromango Island in Vanuatu belong here; this was confirmed from specimens at
K formerly identified as P. insularis. Some specimens from the Solomon Islands now assigned to
P. orarius were also identified as belonging to P. insularis. It seems clear that a critical revision of
that species is also needed, so the maps of both species here presented need to be considered as
more or less provisional.
SOUTH AMERICA

Geography

South America is the world’s third largest continent with an area of 17.85 million km². It has
a roughly triangular shape, with the widest part in the southern tropics stretching 5,200 km
and tapering continuously southward to Cape Horn at 56° S. Its northenmost cape is Punta
Gallinas in Colombia at 12° 20’ N, a distance of ca. 7,500 km. Therefore, the greatest landmass
of the continent is in the tropics and the southern warm temperate to subantarctic climate
zones are on an increasingly smaller landmass with stronger maritime influence. Two major
topographical features dominate the continent: the Andes mountain chain and the Amazon
River, by their sheer size as well as their geographical position. The Andes is the longest con-
tinuous mountain range in the world, over a length of 8,050 km forming a high barrier along
the western margin of the continent. After Asia, South America has the highest mountains in
the world, with Aconcagua, a volcano, topping 6,959 m and more than 50 other summits higher
than 6,100 m. The Amazon and its tributaries form the largest river basin in the world, draining
much of the wider part of the continent from the Andes to the Atlantic Ocean. The Amazon
Basin has filled with sediments from the Andes and other high parts of the continent and is
largely low lying and flat. Other topographical features relevant to conifer distribution are the
Guayana Shield with its table mountains or tepuis in the N and the Brazilian Shield south of
the Amazon Basin. Both are much older than the Andes and heavily eroded. South America
had been the western part of Gondwana from the Jurassic when it started to drift away to the
west after rifting from Africa in the Early Cretaceous. Pushing over the Pacific Nazca Plate it
throws up the Andes, of which the oldest sections are in the far S and the youngest in the N
beginning less than 10 Ma; the uplift still continues over much of its length. The continent was
severed from Antarctica during the Oligocene and connected with and separated from North
America several times in the Tertiary; the present Isthmus of Panama dates from the Pliocene,
ca. 3.1 Ma. The presence and absence of conifer taxa in South America are strongly influenced
by these plate tectonic events.
A major climatic feature with effects on conifer distribution in the Andes is the shift from east-
erly moisture-laden winds in the tropics to westerly currents at around 30° S. From just S of the
equator to that latitude, the W-side of the Andes is the driest, culminating in the Atacama Desert
with near zero precipitation. From ca. 33° S the situation is reversed and the E-side of the Andes
is the driest, but here the effects of both increasing latitude and narrowing of the continent ame-
liorate the contrast. An arid zone known as the South American Arid Diagonal (SAAD) traverses
the S part of the continent from the Atacama Desert to S Patagonia, crossing the Andes in the area
where this climatic shift occurs. In the far south it is wet everywhere all year round. All year high
 south america 429

precipitation also applies in the tropical mountains of N Colombia and Venezuela; in contrast
the highlands of S and SE Brazil have a strongly seasonal climate with extensive dry periods. The
wettest part here is the narrow strip of the Atlantic Rainforest, in which few conifers occur.

Conifers in South America

Families Genera Species Species + infra Countries with Area of occupancy Endemic taxa
conifers in km²
3/8 9/70 34/615 35/794 9/11 15,100/17,850,420 32
Numbers after ‘/’ are global or continent totals.

As can be seen from the table, South America has a low diversity of conifers with only 34 species
(the same number as the larger continent Africa). With only one additional variety it even falls
below Africa in the total number of taxa below the rank of genus. They belong to three families
and nine genera, with the Podocarpaceae by far the more diverse family. There are two species
of Araucaria (Araucariaceae) and three species in Cupressaceae: Austrocedrus chilensis, Fitzroya
cupressoides and Pilgerodendron uviferum. The latter three species are the only ones in their gen-
era and occur only in the southernmost part of the continent. The two species of Araucaria are
also endemic to South America and as can be seen from the table, endemism is very high at
91% for all conifers. Only three species also occur in Central America: Podocarpus guatemalensis,
P. magnifolius and P. oleifolius. All three belong to subgenus Podocarpus with a predominantly
Southern Hemisphere distribution. Perhaps more significantly, no Laurasian taxa, such as the
family Pinaceae and the genus Taxus, have migrated to South America along the Isthmus of Pan-
ama or from the Caribbean Islands. The three genera in Cupressaceae have southern relatives
and so does Araucaria, but it has to be remembered that the latter genus had a more world-wide
distribution in the Mesozoic.
The distribution of conifer species in South America shows a distinct pattern. Most obvious is
the Andean mountain chain, which is followed by conifers for almost its entire length, but with
a substantial gap between 30° and 33° S. A closer look also reveals that conifers tend to occur on
the E side of the Andes in the great arc from Venezuela/Colombia to N Argentina and on the W
side south of the gap, until this predominance on one side or the other of the Andes fades away in
Patagonia in the far south. This is clearly influenced by the pattern of rain distribution mentioned
above. The reason for the gap is probably due to two factors: lack of sufficient precipitation (it
coincides with the SAAD) but also lack of taxa that could adapt or were ‘pre-adapted’ to these
conditions, as exist elsewhere e.g. in California and Mexico. This section of the Andes has some
of its highest peaks but also its driest parts, with on average between 400 and 600 mm annual
precipitation, much of its as snow. With most of the species being podocarps and at least four
of the other five conifer species present in South America also requiring ample moisture, the
genetic potential for adaptation to dryness may not have been available. The SAAD is estimated
to have originated 14–15 Ma and indeed no conifer species cross it, while most lineages to the
S are of older Gondwanan origin. This pattern has been recognised in other plant groups, e.g.
Alstroemeriaceae (Chacón et al., 2012). Lack of precipitation also accounts for the large empty
space in Argentina and Uruguay with mostly steppes (pampas) and semi-deserts to deserts in
the rain shadow of the Andes. On this E-side of the continent conifers reach their southern limit
at 31° 30’ S, at similar latitude as the conifer gap in the Andes. Conifers are widely distributed in
S Brazil and become more scattered further N in the Cerrado, a region of seasonally dry uplifted
and eroded plateaus with a savanna-type vegetation, where conifers tend to follow streams. They
avoid the Atlantic coast, where the Atlantic Rainforest (Mata Atlántica) has no conifers except
in its southernmost part S of Rio de Janeiro. North of the Amazon, conifers are scattered on the
Guayana Shield in extreme E Colombia, Venezuela, NE Brazil and Guyana. Some species of Podo-
carpus are (near) endemic to the tepuis in this area. The great void in conifer distribution is the
430 south america

Map SA-1. Distribution of conifer species in South America.

Amazon Basin. Except for a few localities in its westernmost part, near the Andes, no conifers
are known to occur naturally in this enormous expanse (Dalling et al. in Turner & Cernusak, eds.
2011). It is a similar situation as in the Congo Basin of Africa (see p. 459) and it must have similar
causes. Interestingly, there is some palynological evidence for the presence of podocarps in the
Amazon Basin during glacial maxima of the last Ice Ages. Global cooling may have driven coni-
fers from the Andes into the basin, where the tropical lowland rainforest was possibly reduced
to refugia (Haffner, 1969; Prance, 1982). The refugia hypothesis has been opposed more recently
on the basis of palynological and ecological evidence, but it seems uncontested that the Amazon
Basin experienced a cooler climate during Pleistocene glacial maxima.
Because South America shows these distinct patterns of general conifer distribution and spe-
cies faithfully follow these patterns with few exceptions, we discuss the biogeography of conifers
by the regions in which they occur.
 south america 431

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Map SA-2. Map of South America showing num-
bers of conifer genera per one degree cell. The areas
with the highest numbers of different genera (5–6
genera per degree cell) are in Biobio and La Arau-
cania (Regions VIII and IX) in Chile.
Map SA-3. Map of South America showing num-
bers of conifer species per one degree cell. The
areas with the highest number of different species
(7 species per degree cell) are again in Chile but
also in La Paz, Bolivia and in S Ecuador.

Map SA-4. Rarity values per one degree cell. Cell score is the total of the rarity calculations for each species
in the cell. The rarity calculation is 1 divided by the total of cells occupied by a species. Thus, a species that
occurs in one cell scores 1 while a species that occurs in 10 cells scores only 0.1.
432 south america

The diversity maps SA-2 ans SA-3 show a concentration of diverse 1 degree cells in Chile, espe-
cially when calculated at the genus level. Chile has a number of conifer genera that are absent in
the rest of South America. However, several of these genera are monospecific while Podocarpus is
more diverse, even in South America. Therefore the score in some Andean degree cells at species
level becomes as high as in Chile. Overall, the modest taxonomic diversity of conifers in South
America is situated in the Andes. In map SA-4 the distribution of rare species, i.e. the narrow
endemics, is quite different from the diversity maps. Chile no longer scores highest, instead local
endemism is found in approximately the same degree cells as species diversity in the Andes N of
the gap at 30–33° S but also in other regions. Most notable are the tepuis mountains of the Guy-
ana Shield, where some species of Podocarpus are limited to a few of these table mountains. The
single red cell in Rondônia, Brazil, indicating narrow endemism, is perhaps an artifact of limited
collecting (see also map SA-28 on p. 447).

The Andes North of 30° South

This part of the Andes is situated from N to S in Venezuela, Colombia, Equador, Peru, Bolivia and
Argentina. In the northernmost section of the mountain chain, the range divides into three ranges
(cordilleras) in Colombia, lying more or less parallel NNE to SSW. The Cordillera Oriental divides
once more near the border with Venezuela, with the Cordillera de Merida oriented to the NE into
Venezuela and terminating the Andes near Caracas on the Caribbean coast. The Andes chain is
at its narrowest in Ecuador, more or less forming a single range of mountains. Southeastward in
Peru and Bolivia it widens, with parallel ranges separated by high altitude valleys or plains (Alti-
plano) until the chain again turns N/S and then narrows at around 30° S where the ‘conifer gap’
begins. All species present in this part of the Andes belong to the Podocarpaceae, in three genera
Podocarpus, Prumnopitys and Retrophyllum. Ten species are endemic (or nearly endemic) to the
Andes, four species also occur elsewhere but have major distributions in the Andes, and one spe-
cies, Podocarpus tepuiensis occurs mainly elsewhere. The latter species is mapped and discussed
below in the Guayana Shield section and P. guatemalensis is treated in the chapter Mexico, Cen-
tral America and the West Indies, as its main distribution is in Central America.

Prumnopitys

The genus Prumnopitys has nine species, distributed disjunctly in Central and South America,
New Zealand, New Caledonia and Queensland (Australia). One species occurs in Central America,
four in the Andes, two in New Zealand, and one each in New Caledonia and Queensland. Three
species are endemic to the part of the Andes here discussed, so the biogeography of the entire
genus should be discussed here first. This distribution provides another example of Gondwanan
vicariance, but with possible dispersal at regional level.

Map SA-5. The distribution of the genus Prumnopitys on two sides of the southern Pacific Ocean.
 south america 433

Objectively, it is possible to view this pattern of distribution from two different geographical perspec-
tives. Either the genus Prumnopitys occurs on two opposite shores of the southern Pacific Ocean, or
its disjunction has nothing to do with that intervening ocean but with Antarctica, which lies over
the South Pole between New Zealand and the southern tip of South America. Which of the two
connecting routes is more likely to be the one that actually played a role in its present distribution?
Three approaches to this problem can be pursued: evidence from plate tectonics and continental
drift, evidence from the fossil record and phylogenetic inference from molecular data. Reconstruc-
tions of past positions of land masses could indicate land connections that no longer exist between
the disjunct populations. Obviously no fossils of land plants are expected to be found in the ocean,
but that does not disprove the possibility of dispersal across it. Phylogenetic inference could indicate
which of the nine extant species is most basal in the tree, suggesting an area of origin of the genus
and subsequent spread of other taxa by whatever means. The best approach would combine results
from all three disciplines to see if there is congruence between these lines of evidence. The land-
masses and islands where Prumnopitys is present today are all parts of the former supercontinent
Gondwana. The fossil record indicates presence of the genus in New Zealand (Palaeocene) and in
Tasmania (Eocene) (Hill & Brodribb, 1999). The phylogeny reconstruction is inconclusive. In the
analysis by Biffin et al. (in Turner & Cernusak, eds. 2011) of Podocarpaceae, the most comprehen-
sive of its kind to date, five species of Prumnopitys form a clade with Sundacarpus amarus and this
clade further divides in two clades. One clade unites Sundacarpus amarus (Malesia/Queensland)
with P. andina (N Andes) and P. taxifolia (New Zealand); the other clade has P. ladei (Queensland)
with P. ferruginea (NZ) and P. ferruginoides (New Caledonia). This result cannot tell us where the
ancestor of the genus might have been. Perhaps adding the four missing species would change the
topology of the tree, but this requires additional research. As is the case with other genera that
display similar distributions with extant and/or fossil taxa (Acmopyle, Athrotaxis, Dacrycarpus, Lepi-
dothamnus and Retrophyllum, for the latter genus see map AU-100 on p. 410) the South America/
SW Pacific connection is best interpreted as cases of disjunction caused by extinction in Antarctica
(mainly) and thus as a good example of vicariance biogeography involving a Gondwanan origin.

Map SA-6. Prumnopitys harmsiana [n=29, m=29,

h=10, 1854–2004]

Prumnopitys harmsiana is the most wide-

spread of the three species in this part of the
Andes. It occurs from N Venezuela (Cerro
Avila near Caracas) along the Andes as far
S as central Bolivia. A disjunct population is
known in the Sierra Nevada de Santa Mar-
tha in N Colombia, an isolated massif that
rises to 5775 m. Other gaps apparent on
the map, e.g. the section of the Andes in S
Colombia and Ecuador, are possibly an arti-
fact of the paucity of herbarium collections.
Due to identification problems only verified
specimens have been used for mapping; the
range of this species partly overlaps with
that of P. montana. Prumnopitys harmsiana
is a relatively uncommon tall tree to 35 m
occurring in (lower) montane rainforest
between 1000 m and 2200 m altitude. Because of its size and useful timber this species has been
targeted by loggers and its scarcity may also be an effect of this, especially in heavily deforested
countries such as Ecuador.
434 south america

Map SA-7. Prumnopitys montana [n=43, m=42, h=8,

1845–2005]

Prumnopitys montana is relatively common in

the Andes from Venezuela (Cordillera de Mer-
ida) to central Peru (Junín). In Ecuador, herbar-
ium collections are all from the southern part
of the country, so a gap is apparent from there
northward across the border with Colombia.
Deforestation is probably partly responsible for
this distribution pattern, but since herbarium
collections represent historical occurrences (in
Ecuador for this species from [1847] 1945–1998
in the Conifer Database from which this map is
drawn) it is unlikely to be the only cause. Also,
other conifers have been found to occur in this
gap. Much of the deforestation occurred in the
last few decades. Prumnopitys montana has an altitude range from 1500 to 3600 m and so it occurs
from montane forest to high montane cloud forest; at the maximum altitude it is often a stunted
tree but on lower slopes it can reach 30 m. While larger trees are equally valued for timber as the
previous species, the trees on high ridges often escape logging as they are either inaccessible or
too small to take. Many large trees can still be found in former forest turned to pasture as such
trees were left standing to give shade to cattle. As regeneration in grazed land is unlikely, in due
course the species will likely disappear from these lands unless it is planted. It is listed as Vulner-
able (VU) on the IUCN Red List.

Map SA-8. Prumnopitys exigua [n=27, m=26, h=7,

1951–2007]

Prumnopitys exigua has a very limited distribu-

tion in central Bolivia, in the departments of
Cochabamba and Santa Cruz, spread mainly
in the Cordillera Oriental but also in part of
the Cordillera Central. Prumnopitys exigua is
a montane to high montane tree of moderate
size, ocurring in evergreen cloud forest at alti-
tudes between 1850 m and 3000 m. Its habitat
is similar to that of the widespread species
P. montana, but that species is separated from
P. exigua by a distance of 1000 km. Occurring in the easternmost ranges where the Andes chain
is widest, this species is a good example of the tendency of conifers to occur on the wettest side
of the Andes. A wealth of epiphytic plants growing upon the trees in these forests indicates the
very high, year-round humidity in which P. exigua thrives. Deforestation to expand agriculture
(pasture) is affecting the species at the lower elevations and it could become threatened with
extinction in future if this trend continues.

Retrophyllum

The genus Retrophyllum has five species, distributed in Malesia, the SW Pacific and South America,
mapped (No. AU-100) and discussed on p. 410. The single species in South America has a simi-
lar distribution as Prumnopitys montana, from W Venezuela (Cordillera de Merida) to NW Bolivia
 south america 435

Map SA-9. Retrophyllum rospigliosii [n=25, m=25,

h=8, 1911–2007]

(La Paz), but it is found in disjunct locations

along the Andes chain. There are apparent con-
centrations in the Cordillera de Merida, in the
far S of Ecuador (Cordillera del Condor, Parque
Nacional Podocarpus), central Peru (Pasco,
Junín) and in NW Bolivia (La Paz). Scattered
collections were made in Colombia and N Ecua-
dor. These regions are well collected botanically
and should have yielded more if there was more.
Similarly, although effects of collecting bias can
not be ruled out, the concentrations showing on
the map are likely to reflect a distribution pat-
tern that is real. This is demonstrated when all
conifers are mapped (see map SA-1 on p. 430);
the gaps are then sparsely filled with occur-
rences. A few species occur there in low densi-
ties but many are absent. Maps published by
Dalling et al. (in Turner & Cernusak, eds. 2011) for Neotropical species of Podocarpus show similar
patterns in the Andes (they are based on much the same data). The “multiple disjunct populations”
observed for most species are said to be “suggestive of habitat specialization or refugial distribu-
tions”. The few data that are available point at habitats well known for better studied conifers in
tropical forests: they tend to occupy acidic, leached soils, ultramafic substrates and mountain ridges
and summits with little or no fertile soil. Whether this can explain the large gaps in the distribu-
tion of Retrophyllum rospigliosii remains doubtful. The altitudinal range of this species is from 1500
to 3300 m (-3750 m in Colombia and Peru) so it occurs in wet rainforest up to high altitude cloud
forest or ‘mossy forest’. It can form more or less extensive pure stands on exposed sites, but is more
often found scattered among angiosperms or sometimes with Prumnopitys spp. As a tall tree (to
45 m) at lower elevation it is a target for logging which is often followed by deforestation. Retrophyl-
lum rospigliosii is listed as Vulnerable (VU) on the IUCN Red List.

Podocarpus

Map SA-10. Podocarpus glomeratus [n=36, m=34,

h=9, 1923–2007]

Podocarpus glomeratus is a shrub or a stunted

tree to 12 m tall occurring in high montane to
subalpine forests and woodland or scrub, usu-
ally between 2500 and 3600 m. It is distributed
from Ecuador to Bolivia and shows a pattern
of disjunct occurrences similar to other species
of Podocarpaceae in the central and northern
Andes. In Peru, it is known from only two loca-
tions, the northern one in the Sierra de Chaglla
in Huánuco and the southern one in Abancay
in Apurímac. Both are situated in central ranges
of the Andes chain. More extensive distributions
are present in both Ecuador to the north and in
Bolivia to the south of these isolated occurrences.
At such distances of separation, edaphic factors seem to be of minor relevance to the observed
436 south america

­ attern, as similar conditions would exist in the intervening gap regions. During glacial periods of
p
the Pleistocene, the central ranges of the Andes were heavily glaciated and forest was driven to low
elevations or went extinct. Perhaps the two Peruvian populations are relicts of refugia.

Map SA-11. Podocarpus parlatorei [n=44, m=42, h=10, Podocarpus parlatorei in secondary vegetation in
1846–2008] Tucumán, N Argentina. Photo credit J. F. Pensiero.

Podocarpus parlatorei has a nearly continuous distribution along the eastern side of the Andes from
central Bolivia (Cochabamba and Santa Cruz) to northern Argentina (Tucumán), a distance of ca.
1200 km. No other species in the Andes N of 30° S display such continuity. Records for Peru posted
in GBIF on the Internet (e.g. in the Sierra de Chaglla) are misidentified. They were included in maps
generated from GBIF data published by Dalling et al. (in Turner & Cernusak, eds. 2011) although their
text is correct in stating that P. parlatorei only extends “north into Bolivia”. As we have explained in
the chapter Creating the Atlas of the World’s Conifers, including GBIF or similar source data without
scrutiny leads to errors. Podocarpus parlatorei is usually a shrub or stunted tree at high altitudes
between 2000 and 3150 m, but can become a moderately tall tree to 20 m at lower elevation, where
it is sometimes abundant on old river terraces and on river banks. It is a pioneer species with a key
role in the forest dynamics. Its seeds or ‘fruits’ are a food resource for endangered and of restricted
range bird species, such as Penelope dabbenei (Cracidae) and Amazona tucumana (Psittacidae).

Map SA-12. Podocarpus rusbyi [n=24, m=23, h=10,

1886–2005]

Podocarpus rusbyi is almost entirely confined

to the Andes in Bolivia, but with one locality in
Peru (Machu Pichu in Cusco). The localities are
in the Cordillera Oriental of the main Andes
chain and in the more eastern and lower moun-
tain range of the same name in Bolivia. A col-
lection (R. B. Foster et al. 10934 with duplicates
at K end NY) from the Cerro de Pantiacolla in
Peru (“700–1300 m”) has been identified as this
species, but belongs to Podocarpus oleifolius.
 south america 437

The altitude range of P. rusbyi is 1500–3350 m, with few collections made below 2000 m. It occu-
pies similar montane to high montane habitats as P. parlatorei within the same altitudinal range,
but in Bolivia the two species are geographically separated. It is commonly associated in the for-
est with Meliaceae, Myrtaceae and other angiosperm trees; at high altitudes it forms thickets with
other dwarfed trees and shrubs. This species is listed as Vulnerable (VU) in the IUCN Red List.

Map SA-13. Podocarpus sprucei [n=17,

m=16, h=7, 1858–2005]

Podocarpus sprucei occurs in the

Andes of Ecuador and adjacent
parts of Peru. It is a species of
high montane to subalpine for-
est, found at altitudes between
1800 and 3900 m. A tree at the
lower altitudes attaining 20 m, it
becomes dwarfed at or just below
the tree line, where the forest gives
way to páramo, a treeless vegeta-
tion often dominated by tussock
grasses and with conspicuous
presence of cushion plants and
rosette-forming large herbs. Log-
ging of larger trees but especially
deforestation and land conversion
to pasture have caused serious
decline of this species, which is now listed as Endangered (EN) on the IUCN Red List.

Map SA-14. Podocarpus salicifolius [n=17, m=16, h=9, Podocarpus salicifolius. A sapling tree grow-
1843–2000] ing in a gully in secondary forest, Bolivia.
Photo credit Graham Charles.
438 south america

Podocarpus salicifolius has its main distribution in the mountains W of Caracas, Venezuala, the
most northeasterly extension of the Andes chain along the Caribbean coast. It is also reported
from a few disjunct localities in Colombia, NW Brazil, Peru and Bolivia, some of which have not
been mapped for lack of verifiable herbarium specimen data. It is unlikely to be as rare in parts
of the Andes outside Venezuela as these data suggest; the species is probably under-collected
despite its early description and naming in 1847. Podocarpus salicifolius occurs in the Andes
ranges in montane to upper montane rain forest or cloud forest, at altitudes between 1200 m and
2150 m a.s.l.

Map SA-15. Podocarpus pendulifolius [n=7,

m=7, h=3, 1944–1988]

This species has a limited distribution

and occurs in the Cordillera de Mer-
ida, the extension of the Andes in W
Venezuela. It is known from two high
altitude areas: Páramo de la Cristalina
and Páramo de la Negra, at a distance
of 100 km from each other. It does not
actually occur in the páramo vegeta-
tion zone, but below it in forests at
altitudes between 1400 and 3000 m. At
the highest elevation this forest is low
and merges via a scrubland ecotone
to the subalpine and alpine páramo
dominated by grasses and tall herbs.
Podocarpus pendulifolius is most often
seen in the high transition zone and less frequently in the taller forest below, where it can attain
20 m. This rarity is partly the result of logging and forest clearing and this species is listed as
Endangered (EN) on the IUCN Red List.

Map SA-16. Podocarpus celatus [n=27,

m=25, h=7, 1866–2007]

Podocarpus celatus occurs scattered in

the Andes from Colombia and Ecua-
dor to Bolivia and also in the Guayana
Highlands of Venezuela. Another well
known and well collected location is
Maynas in Loreto, Peru, which is in
the Peruvian Amazon at only 130–140
m a.s.l. The highest herbarium collec-
tion record is at 1930 m in the Cordil-
lera de Nanguipa of Ecuador. Some
very disjunct localities are in Brazil, in
Mato Grosso and in Goiás (the latter
is mapped here, collected in 1971
at 1250 m in cerrado on sandstone
outcrops in Chapada dos Veadeiros
National Park). This is an erratic dis-
tribution spread over an enormous region and as such difficult to interpret. Soil types mentioned
are laterite, sandstone outcrops and white sand, i.e. nutrient-deficient substrates. Not uncommon
 south america 439

terrain for podocarps to retreat to if faced with competition, and plenty of it is available in this vast
land. Therefore the apparent rarity is not due to scarcity of special habitats. More likely is that it is
vastly under-collected. The interior of Brazil in particular remains poorly inventoried botanically,
with collecting efforts concentrated in scattered centres (Nelson et al., 1990).

Map SA-17. Podocarpus magnifolius [n=20, m=20,

h=7, 1939–2006]

This species is known from widely separate

locations in the Andes, from E and S Venezuela
and from Panama. In the Andes, it occurs from
near Caracas, Venezuela to La Paz, Bolivia, but
with apparent great gaps, e.g. missing in a long
section from S Colombia through Ecuador and
N Peru. In Panama it is only know from the E
half of the country. While the occurrence in E
Venezuela is in the Venezuelan Highlands, the
locality in the S of the country is the Pico de
Neblina, where it may occur on both sides of
the border with Brazil. While generally giving
the extent of occurrence (EOO) this distribu-
tion pattern seems to indicate limited avail-
able data, i.e. herbarium collections, as has
been concluded for some other species in this
region. Podocarpus magnifolius is a forest tree
to 30 m tall with leaves that measure up to 29 cm long and 3 cm wide, but can be as small as 3
cm long and 12 mm wide; much depending on tree age and exposure to sunlight. Large leaves will
enable the species to be shade-tolerant and compete with large-leaved angiosperms. The altitude
range is considerable, from 850 to 2900 m and this too has a bearing on leaf size, with smaller leaves
found at highest altitudes. Little is known about the (possible) habitats this species can occupy,
which may be a reason why it has not been found often.

Map SA-18. Podocarpus oleifolius [n=104,

m=97, h=16, 1844–2009]

Podocarpus oleifolius is widespread in

the Andes chain from Venezuela to
Bolivia, with few gaps except possibly
one in Peru between Pasco and the
southern part of Cuzco, a section of
the Cordillera Oriental from where we
have found no herbarium records. It is
also known, more scattered, from sev-
eral places outside the Andes in Ven-
ezuela, Colombia and Bolivia, and it
extends into Central America as far N as
Veracruz in southern Mexico. There an
apparent gap is in Panama, where it is
recorded from near the Colombian bor-
der and from mountains in the western
part of the country, but not in between.
440 south america

This is the largest range of any species of conifer in South America and one of the largest in the
world for a species in the Podocarpaceae. Podocarpus oleifolius is a montane to high montane
species occurring in primary semi-deciduous and evergreen forests. Commonly a tall tree in the
forest, it mixes with numerous angiosperms and ocasionally other conifers, especially podocarps.
Its altitudinal range is large, from 700 to 3450 m, but most commonly its is found above 2000 m;
at the highest elevations it is a common component of stunted cloud forest rich in epiphytes,
especially bromeliads and orchids.

The Guayana Shield

The Guayana Shield is one of the oldest parts of South America, with rocks dating back to the
late Precambrian 800–600 Ma. This ancient continental core extends from E Colombia across
Venezuela and N Brazil to the Guyanas. It is heavily eroded and the general elevation is low,
around 500 m, but in several places much higher plateaus, mostly reduced to table mountains or
tepuis, rise to 1500–2500 m a.s.l. The highest is the Pico de Neblina on the Brazilian-Venezuelan
border at 3014 m, deeply dissected on all sides and the highest point in South America outside the
Andes. Most tepuis are in the E part of the Venezuelan Guayana Highlands, where 115 have been
counted. Above a sloping apron of talus they typically consist of massive blocks of Precambrian
quartz arenite sandstone with perpendicular sides (faces) and flat but eroded and dissected tops,
often dotted with large sinkholes. Although some tepuis occur in random groups, many are very
isolated and because they are ancient ‘islands in the sky’ they have led to substantial endemism.
Three species of Podocarpus (Podocarpaceae) are mainly distributed on the Guayana Shield, one
species, P. roraimae is endemic, one species also occurs in N Venezuela and the third species is
also found in Ecuador.

Map SA-19. Podocarpus roraimae

[n=14, m=14, h=5, 1909–1988]

Podocarpus roraimae was des­

cribed from Mt. Roraima, a
famous tepui on the border
between Guyana, Venezuela
and Brazil (the three-country
point is on its flat top at 2810
m) and has subsequently been
collected from several tepuis
in Venezuela, including the
Neblina Massif (Pico de Neblina)
in the south. The altitude range
recorded is 1800–2700 m and it
is virtually limited to the tepui
summits. This species occurs in
small stands of shrubs and trees
in a mozaic vegetation of ombrotrophic bogs dominated by grasses and herbs and interspersed
low scrub and woodland. It can sometimes be found in ravines leading to the edge of the tepuis,
where forest can be somewhat higher and it may attain tree size. The tepuis summits are defi-
cient of nutrients (there are thus many carnivorous plants) but very wet and often shrouded
in clouds.
 south america 441

Map SA-20. Podocarpus steyermarkii [n=13,

m=13, h=5, 1944–2003]

Julian A. Steyermark (1909–1988) was

the great botanist, plant collector and
describer of the flora of the tepuis and
this species was named after him. The
main distribution is on the tepuis of
Guyana (Pakaraima Mts.) and Venezuela
(Carrao-tepui, Uaipan-tepui, Cerro Jaua
and Pico de Neblina), but two collections
in the Conifer Database are from the Cor-
dillera de Merida and its extension near
Caracas in N Venezuela. Not many col-
lections are known and this may be the
rarest of the three species here mapped.
The occurence in the Andes of N Ven-
ezuela strongly hints at relict popula-
tions perhaps dating back to the glacial periods of the Pleistocene, when a cooler climate in the
intervening lowlands may have allowed this conifer to spread northwards. This species has now
been found in cloud forest with a canopy to 20–25 m tall, as well as on the tepuis summits in
low herbaceous or scrub vegetation in areas with blanket bogs with grasses, ferns, pitcher plants,
orchids, mosses, and low scattered shrubs to 1.5 m tall, interspersed with thickets and dwarf forest
including palms. It has been found associated with the other two species of Podocarpus of the
tepuis in a few localities.

Map SA-21. Podocarpus tepuiensis

[n=20, m=20, h=6, 1944–2006]

Podocarpus tepuiensis has its

main distribution area in the
Guayana Highlands of Ven-
ezuela and Guyana, where it
occurs mainly on the tepuis
of the Guayana Shield, which
are sandstone plateaus. Other
localities in S Venezuela (Ama-
zonas) are in the lowlands. A
second centre of occurrence
is known since 1993 in Ecua-
dor, where it occurs in the
Cordillera del Condor and in
the Cordillera Sacha Llangan-
ates. In Venezuela Podocarpus tepuiensis occurs from 100 to 2450 m; in Ecuador from 1600 to 2740
m. In Venezuela, off the tepuis, it is mainly found along streams in gallery forest, on the tepuis
in dwarf forest patches. The lowland localities are on white sand at the edge of savanna and for-
est. In the Andes of Ecuador the habitat of this species is most often outcrops of quarzite amid
sandstone formations similar to the Guayana Shield mesa sandstones, but in the Cordillera Sacha
Llanganates it was found on a granite ridge. On the tepuis, it may sometimes be associated with
P. roraimae and P. steyermarkii, but these two podocarps do not occur at lower altitudes than
1800 m. The wide altitudinal range, the two very disjunct centres of occurrence and the relatively
recent discovery in Ecuador together suggest that its range is incompletely known at present.
442 south america

Brazil

Brazil is the largest country in South America with an area of 8.5 million km². In the north by far
the largest topographical feature is the Amazon Basin, much of it below 200 m and in the southern
parts rising to 300–500 m a.s.l. Only on the border with Venezuela and W Guyana is there any
high elevated terrain, with the Serra Imeri/Neblina Massif, the Serra Parima and the Pakaraima
Mountains. South of the Amazon Basin the land generally rises, with hills in Rondonia and Mato
Grosso and low mountains in the Planalto of Goiás and Bahía to about 1000 m. The climate is
predominantly tropical. In the southern part of the country there are more low mountains, eroded
from the ancient Brazilian Shield to altitudes mostly not exceeding 2000 m. The Amazon rainfor-
est which still covers much of the Amazon Basin gives way in the south to the Cerrado, a vast
tropical savanna region covering 21% of the land area of Brazil. Seasonality is pronounced, with
a long dry season in the southern winter. The most relevant aspect of the vegetation structure to
conifers is that tall trees are mostly restricted to strips of gallery forest following streams, with ‘wet
campo’ forming an ecotone between these and the dry savanna proper. In the SW of Brazil there
are vast swamps in Mato Grosso. Along the Atlantic coast from Bahía to Rio Grande do Sul occur
the remnants of a previously nearly continuous Atlantic rainforest, distinct biologically from the
Amazon rainforest. The far S of Brazil from São Paulo to Rio Grande do Sul has a subtropical to
warm temperate climate and the natural vegetation is a mosaic of forest and grassland, character-
ized by the conifer Araucaria angustifolia. This is also the most densely populated region of Brazil
and much of the vegetation has been altered or removed to make way for food or tree crops.

Map SA-22. The distribution of conifer species in Brazil.

 south america 443

Brazil is for its size poor in conifer diversity, with only nine species + one variety occurring within
its borders. All except Araucaria angustifolia (Araucariaceae) belong to the family Podocarpaceae.
The Amazon Basin is devoid of conifers (plantations are here excluded) and we have discussed
the possible causes at the beginning of this chapter. In the Cerrado conifers are most common
in the higher elevation areas of Goiás, Bahía and Minas Gerais, elsewhere they are rare, as is
also the case on the northern and southern margins of the Amazon Basin in the western part of
Brazil. Conifer species are most numerous and occur most abundantly in the southernmost part
of Brazil, but they also reach their southern limit there, as can be seen on map. SA-1 on p. 430,
showing conifer distribution in the whole of South America. Of the nine species native to Brazil,
Podocarpus celatus is more widespread in the Andes and has been mapped and discussed in that
section (map SA-16 on p. 438).

Araucaria

Map SA-23. Araucaria angustifolia

[n=40, m=32, h=20, 1817–1997]

Araucaria angustifolia is one

of two species native to South
America (for the genus see map
AU-74 on p. 391) and it is nearly
endemic to Brazil, with just a
small part of its global popu-
lation in Misiones, the narrow
strip of Argentinian territory
between Brazil and Paraguay,
crossing the border into Para-
guay at one or two localities.
Its extent of occurrence (EOO)
is from near Belo Horizonte
in Minas Gerais to central Rio
Grande do Sul, a distance of ca.
1500 km, and from the Serra do Mar to the Paraguay border with Argentina, nearly 500 km.
Within this large area its present area of occupancy (AOO) is but a fraction of what it was
before European settlement, having been reduced from ca. 200,000 km² to a mere 4000 km²
(Koch & Celeste Correa, 2002). What remains are fragments of the araucaria forests, some quite
large and in protected areas, others mostly small or very small and on private land. Therefore,
the map still records the EOO, or most of it, as it was 200 years ago. Araucaria angustifolia is
listed as Critically Endangered (CR) on the IUCN Red List despite this and the protected areas,
because the decline has not stopped and (illegal) logging and replacement by plantation of
exotic trees is still ongoing. The natural vegetation consists of a mosaic of araucaria-dominated
tall forest (capâo) amid extensive grassland (campo limpo), somewhat similar to the habitat
of Araucaria bidwillii in Queensland, Australia (map AU-9 on p. 349). The altitude range for A.
angustifolia is (300–)500–1800 m. Its northern limit coincides with the southern margins of the
Cerrado where a dry season reduces annual precipitation. In the south the araucaria forests
are replaced by the grasslands of the pampas which commence in the valleys of the Jacuí and
Ibicuí Rivers in Rio Grande do Sul. It is therefore primarily precipitation that is the limiting
factor in its natural distribution.
444 south america

Araucaria angustifolia in a natural forest remnant between Cambara do Sul and Tainhas in Rio Grande
do Sul, Brazil. The araucarias are emergents above a lower canopy of angiosperms and occasional trees of
Podocarpus lambertii. Photo credit Aljos Farjon.

Podocarpus and Retrophyllum

Map-SA-24. Podocarpus brasiliensis

[n=34, m=31, h=4, 1909–2009]

Podocarpus brasiliensis occurs S of

the Amazon Basin in Bahía, Dis-
trito Federal, Goiás, Mato Grosso,
Minas Gerais and Rondônia. North
of the Amazon it has sporadically
been collected in Roraima, and
across the border in the Guayana
Highlands of Venezuela. This spe-
cies has quite often been misiden-
tified in herbaria, e.g. as P. sellowii
and a few collections remain of
uncertain identity as they lack
mature and/or fertile material.
Therefore it is suspected that more herbarium specimens of this species exist, awaiting correct
identification. Despite this, it is probably the case that P. brasiliensis is a rare species in Mato
Grosso, Rondônia and Roraima, as well as in Venezuela. In the SE of Brazil this species is mostly
found in the uplands as it is there seasonally drier than in the western and northern parts of the
country. This is Cerrado country, and P. brasiliensis keeps to gallery forests along streams in most
localities. In Venezuela it was found on ridges and spurs of the tepuis in a vegetation that is tran-
sitional between the tepuis top dwarf forest and the tall tropical rainforest below. More collecting
and research are needed to obtain a better understanding of the distribution and ecology of this
apparently widespread species.
 south america 445

Map SA-25. Podocarpus sellowii var. sellowii

(black) [n=31, m=24, h=10, 1874–1995], P. sell-
owii var. angustifolius (red) [n=8, m=6, h=4,
1872–2002]

Podocarpus sellowii occurs in the states of

Paraná, Rio Grande do Sul, Rio de Janeiro,
Santa Catarina and São Paulo. It is mainly
distributed in the coastal Atlantic rain-
forest (Mata Atlantica), but with a few
outlying sub-populations further inland.
The Atlantic rainforest in the S of Brazil is
subtropical evergreen and grows on acidic
soils derived from granite and gneiss.
Podocarpus sellowii is a montane species
growing at altitudes between 800 and 1800 m on relatively dry sites with thin soils where the
forest canopy is more open and generally lower than 20 m, often not more than 10–12 m. In the
northern half of its range these tend to be eastern slopes of the coastal mountains. Near Rio de
Janeiro P. sellowii var. angustifolius occurs in two localities: Serra dos Orgãos and Pico do Tingua.
Much of the Mata Atlantica has disappeared under human pressures causing deforestation and
only scattered fragments remain in this southern part of Brazil. This species is listed as Endan-
gered (EN) on the IUCN Red List, largely due to extreme decline of its habitat in the past. Few of
the localities here mapped are within protected areas.

Map SA-26. Podocarpus lambertii [n=50, m=48,

h=7, 1908–2005]

More widespread than the previous spe-

cies and not limited to coastal mountains,
P. lambertii is found in São Paulo, Rio
de Janeiro, Paraná, Santa Catarina, Rio
Grande do Sul and Minas Gerais in Bra-
zil and in Missiones in NE Argentina. This
species occurs in gallery forest along riv-
ers and in forest margins, especially in the
southern part of its range (Santa Catarina
and Rio Grande do Sul) and elsewhere
mainly in a mosaic of habitats known as
‘campo rupestre’ (‘rocky grassland’). Aside
from taller vegetation following streams,
P. lambertii can occur in rock outcrops,
which give water from run-off and pro-
tection from fires. As can be expected this species usually remains a small tree not taller than
10–12 m, often with a broad, irregular crown and, like most podocarps, is slow growing. In Santa
Catarina and adjacent parts of Rio Grande do Sul P. lambertii is common in the drier type of
Atlantic rainforest on the W-side of the coastal mountains and is there sometimes joined by
P. sellowii. Due to its greater adaptability to adverse conditions P. lambertii is less limited in its
habitat requirements and consequently has a much wider range in southern Brazil than P. sell-
owii. Of the two, P. lambertii has the smallest leaves, only half the size of those of its more coastal
neighbour and a good indicator of its less favourable environment.
446 south america

Podocarpus lambertii in Caracol State Park, Rio Grande do Sul, Brazil. Photo credit Aljos Farjon.

Map SA-27. Podocarpus transiens [n=21,

m=20, h=5, 1909–1997]

Podocarpus transiens is, like P. lambertii,

a small tree with small leaves. It extends
Morro do Chapéu farther N in Brazil and is one of its
few endemic conifers, found in Bahía,
Goiás, Minas Gerais, Paraná and Santa
Catarina. It is sometimes confused with
P. lambertii and so to map it based on
herbarium collections these need to
have been verified by a specialist. Podo-
carpus transiens is more a forest species,
occurring in remnants of forests that
persist naturally at higher elevations
in mountainous locations surrounded
by ‘campo ruprestre’ or in the N of its
range the Cerrado (savanna). The alti-
tude range is 1000–1800 m based on
data from herbarium labels. Its habitat
seems therefore distinct from that of
P. lambertii but remains in fact poorly
known. Its most well-known locality
is the mountain Morro do Chapéu in
Bahía. Podocarpus transiens is rare and
occurs in small numbers in most of its known localities and whilst not a valuable timber tree it is
still exploited for firewood locally. It is listed as Endangered (EN) on the IUCN Red List.
 south america 447

Map SA-28. Podocarpus acuminatus

(black) [n=3, m=3, h=1, 1985–1986],
P. aracensis (red) [n=3, m=3, h=4,
1978–1986] and Retrophyllum piresii
(yellow) n=1, m=1, h=5, 1976]

All three species in this map have

been described and named on the
basis of very little evidence and
two of them can on that basis only
be provisionally accepted as the
morphological differences with
related species may turn out to be
inconclusive if more specimens
could be studied (Farjon, 2010a,
Vol. 2). Retrophyllum piresii was
published in 1983 but is still only
known from the type collection,
gathered in the Serra dos Pacahas
Novos in Rondônia, Brazil. Podo-
carpus aracensis is confirmed to
occur on Serra Araca in Amazo-
nas, Brazil and on Cerro Yaví, Amazonas, Venezuela. The Pico (Cerro) de Neblina (Venezuela)
specimens mentioned in the protologue are of doubtful identity (Farjon, 2010a, Vol. 2, p. 809)
and here not mapped. Podocarpus acuminatus is known from three localities, in Brazil from the
Serra da Neblina (Amazonas) and in Venezuela from the Pico de Neblina (Amazonas) and the SW
Amuri-tepui (Bolivar). The Neblina occurrences belong to a single population on either side of the
international border and show as one on the map presented here. Little is known of the habitat
of Retrophyllum piresii other than that the type collection was made in a forest at 250 m altitude.
The two species of Podocarpus are found on the tepuis and could be limited to these formations.
Clearly, these taxa are poorly known and not recognized by collectors as distinct. Only more (and
better) specimens and field observations, supported by DNA sampling and analysis, can solve
these issues, but they occur in remote places.

The Andes South of 30° South

The parts of Chile and Argentina discussed here are those to the south of 30° S latitude. This is
the narrow part of the continent that is situated in temperate latitudes between 30° and 56° S
and is strongly influenced climatically by the Pacific Ocean and in the far south also by the Atlan-
tic Ocean. The Andes remains the climatic barrier, but moisture here comes from the western
(Pacific) Ocean and the eastern side lies in the rain shadow. This situation becomes gradually
less pronounced further south until it is no longer apparent because both oceans bring moisture
so that there is nearly continuous rain in Tierra del Fuego. To the E of the Andes in the N of this
region are semi-deserts dominated by cacti and scrub merging further south and east with the
pampas, open plains dominated by grasses. On the W side the Atacama Desert merges southward
with a summer-dry zone with a Mediterranean type climate, especially prevalent in the Chilean
Central Valley. Further south this valley breaks up into an archipelago and the climate is cool and
wet. The Andes chain in this region is older (mainly Oligocene) than the central and northern
parts and therefore lower, but numerous volcanos have created new mountains, several exceed-
ing 3000 m and some still active. On both sides of the Andes divide to the south of 39° S numerous
lakes were formed by the very large glaciers of the Ice Ages. Glaciers and ice fields still present
448 south america

today increase in size further south until the Andes becomes too broken up at Punta Arenas to
sustain large ice fields. The Valdivian and Magellanic temperate rainforests in this region are the
most extensive of their kind in the Southern Hemisphere.

Map SA-29. The distribution of conifer species in South America south of 30° S.

As has been observed in the introduction to this chapter, there is a gap in conifer distribution
in the Andes between 30° and 33° S. Conifers become common and diverse from around 35°
S and are certainly less diverse (3 to 1 species) from ca. 44° S. All occur only on the W side of
the continent and mainly on the W side of the Andes divide. This means that the nine species
of this region are mostly concentrated in an area 1200 km long from N to S and 200–300 km
wide. Although few in number, they are of great biogeographical significance. First, all species
are endemic to the region, some with relatively small ranges within the area just outlined.
Second, they belong to three families and eight genera, four of which are monospecific. And
third, most of these have relatives in Tasmania and/or New Zealand, either extant or fossil. The
three species in Cupressaceae, Austrocedrus chilensis, Fitzroya cupressoides and Pilgerodendron
uviferum are the only species in their genus but the first two genera have been found in the
fossil record of Tasmania (Hill & Brodribb, 1999). Araucaria araucana is one of two species in
Section Araucaria, the other being A. angustifolia of Brazil (map SA-23 on p. 443); other ­species
 south america 449

in the Australasia region and New Guinea are more distantly related. The Podocarpaceae are
represented by Lepidothamnus fonkii with two sister species in New Zealand (maps AU-62 and
AU-63 on p. 383), Prumnopitys andina with other species in South America, New Zealand, New
Caledonia and Queensland, Australia (see for genus map SA-5 on p. 432), Saxegothaea con-
spicua (one species only known from this region) and two species of Podocarpus, a pan-tropical
genus. This geographic pattern and taxonomic distinction strongly indicate that these taxa are
ancient relicts of a conifer diversity that was much greater and more widespread in the past,
with links to Gondwana that became severed by continental drift and the refrigeration of Ant-
arctica. The genus Nothofagus (Fagaceae) has a very similar vicariant distribution, supporting
this biogeographical hypothesis.

Map SA-30. Araucaria araucana

(Araucariaceae) [n=48, m=46, h=12,
1854–2004]

Araucaria araucana has a limited

range with most of the global
population in Chile and some
substantial stands across the bor-
der in Argentina. It occurs from
Volcan Antuco in the north to
the shores of Lake Nahuel Huapi
in the south, i.e. between 37°20’ S
and 40°50’ S. Two disjunct popu-
lations occur in the coastal Cor-
dillera Nahuelbuta of Chile. This
is a difficult conifer to make her-
barium specimens of and conse-
quently the map here presented,
although showing the distribution
range generally, remains incom-
plete. Five important areas with
missing specimens are indicated
by grey ovals. The narrow latitudi-
nal range marks the transition between two climate zones: neotropic and antarctic (Golte, 1993),
with summer-dry conditions to the N and cool-wet conditions to the S. In the Southern Hemi-
sphere S-N climate zones are much narrower than their opposites in the Northern Hemisphere,
where this tree thrives as a planted ornamental across far greater latitude. In the Andes, A. arau-
cana is closely assosiated with volcanic deposits of tephra and scoria, so it acts as a colonizer
after volcanic eruptions. In the coastal ranges some rocks are granitic or metamorphic and these
sustain much smaller populations; one at Villa Las Araucarias (in the westernmost oval) was
found to be genetically distinct (Bekessy et al. 2002). The altitudinal range is 600–1800 m. It often
forms nearly pure stands especially at higher altitudes in the Andes, at lower altitudes it is usually
mixed with Nothofagus dombeyi; in the drier Argentinian forests E of the Andean divide Austro-
cedrus chilensis can be found associated with A. araucana. To maintain dominance especially
over angiosperms A. araucana is both a rapid colonizer after volcanic disturbance and capable of
quick crown repairs from epicormic buds after damage. Over the longer geological time span this
conifer may follow shifting volcanic activity along the Andes chain within the limits of climate,
which itself will change. This species is listed as Endangered (EN) on the IUCN Red List.
450 south america

Araucaria araucana on the lava flows and scoria fields of Volcán Llaima, Parque Nacional Conquillio, Chile.
On these newly created slopes the araucaria is a pioneer tree, often together with Nothofagus spp. but
this conifer can persist in older and taller forest dominated by Nothofagus dombeyi when richer soils have
developed and the forest is not destroyed by another eruption. In those forests it can be much taller and
attain 45–50 m. Photo credit Aljos Farjon.

Map SA-31. Austrocedrus chilensis (Cupres-

saceae) [n=96, m=86, h=19, 17??–2011]

Austrocedrus chilensis is the northern-

most conifer in the region, occurring
from 32° 29’ S to 43° 38’ S; in the northern
2/3 of its range exclusively in Chile, in the
southern part predominantly in Argen-
tina. This shift across the Andean divide
from W to E is caused by the increasingly
wet conditions further south in Chile,
so this species moves over to the rain
shadow side of the Andes at ca. 40° S. We
see here on a smaller scale and in a single
species a reversal of the shift from E to W
of the Andes divide observed for conifers
in the continent as a whole (map SA-1 on
p. 430). Both shifts are determined by
levels of precipitation but this species is
exceptional in South America in that it is
adapted to relatively dry conditions. Aus-
trocedrus chilensis is a pioneer tree colo-
nizing disturbed areas e.g. after volcanic
eruptions or land slides. It often occurs in
 south america 451

pure stands or mixed with Nothofagus dombeyi, but also grows associated with Fitzroya cupres-
soides in the coastal ranges and with Nothofagus spp. in the cooler parts of the Andes. Where it
borders the Patagonian steppe it is the only tree species, but does not attain large size there.

Austrocedrus chilensis on a dry sandy slope in the Andes between Rio Caleufeu and Rio Traful, Parque
Nacional Nahuel Huapi, Argentina. Here the mountains are in the rain shadow of the continental divide,
merging with the Patagonian steppe, and Austrocedrus chilensis is the only conifer and forest forming tree.
Photo credit Aljos Farjon.

Map SA-32. Fitzroya cupressoides (Cupres-

saceae) [n=44, m=38, h=13, 1830–2009]

Fitzroya cupressoides, known as ‘alerce’

(‘larch’) in Chile and Argentina, has a lim-
ited distribution; in Chile it is confined to
Los Lagos (Region X) and in Argentina to
adjacent parts of extreme western Neu-
quén, Rio Negro and Chubut. Extensive
logging particularly in Chile followed by
land conversion has reduced this magnifi-
cent tree, the ‘sequoia of South America’
to scattered occurrences now mainly in
reserves, of which many were established
only recently. Most lowland stands have
all but disappeared and the species is
listed as Endangered (EN) on the IUCN
Red List. The herbarium collections rep-
resented on the map date back to 1830
and some dots may indicate locations
452 south america

where F. cupressoides no longer occurs; however, the majority of collections were made after
1945 and about half after 1980. Conversely, not all current stands of this tree are represented in
the herbaria consulted. In Chile this species occurs in the coastal Cordillera, including the island
of Chiloé, a few locations in the Central Valley and in the western volcanic belt of the Andes. In
Argentina it is found in the wetter parts of the Andes chain around the glacial lakes, usually at
their W ends near the border with Chile. Fitzroya cupressoides is a dominant or emergent tree
usually forming stands of variable size (‘alerzales’) scattered in the Valdivian evergreen rainforest.
Several other Chilean conifers are often present in the lower canopy or understorey, as well as
angiosperms. It is present in very wet, water-logged forest with Lepidothamnus fonkii in bogs and
Pilgerodendron uviferum and F. cupressoides occupying slightly higher ground; this habitat type
is common in the coastal Cordillera. In the Andes a more diverse forest type with conifers and
Nothofagus spp. variously dominant is associated with episodal disturbance, naturally from volca-
nism and more recently also from logging operations. Fitzroya cupressoides is one of the longest
living trees in the world, with record ages counted between 3500 and 4000 years.

Map SA-33. Pilgerodendron uviferum

(Cupressaceae) [n=71, m=55, h=15,
1767–2009]

Pilgerodendron uviferum has an

extensive distribution in the
southernmost parts of Chile and
with Lepidothamnus fonkii (map
SA-34 on p. 453) it is one of the
two southernmost conifers in the
world. It extends from the Cor-
dillera de la Costa in Los Lagos
(Prov. Valdivia) at ca. 39° 40’ S to
Ilha Capitan Aracena in Magal-
lanes at 54° 14’ S. A few localities
are across the border in Argen-
tina, but this is mostly a species
of the Chilean Valdivian and Ant-
arctic rainforests. In Los Lagos,
the northern part of its range,
Pilgerodendron uviferum occurs
with Fitzroya cupressoides and
Podocarpus nubigenus but further
south only P. nubigenus accom-
panies it on the edges of bogs in
which Lepidothamnus fonkii forms
extensive creeping mats. In Los
Lagos Pilgerodendron uviferum
has been extensively logged, here
occur(ed) the tallest trees. It is
listed as Vulnerable (VU) on the
IUCN Red List. This and other
conifers such as Araucaria arau-
cana and Fitzroya cupressoides are legally protected in Argentina and Chile and are listed on
Appendix I of CITES, effectively prohibiting the export of their timber. They are also protected
in numerous reserves among which are several national parks. The problem is law enforcement;
under a growing rural population the pressure to convert forest to agricultural uses such as pas-
ture land is ongoing and all three species are still declining, if no longer as fast as in the past.
 south america 453

Pilgerodendron uviferum in a coastal bog in Parque Nacional de Chiloé, Section Anay, Chiloé Island, Chile. In
open marshes on highly acidic soil this tree never grows taller than a few meters. Photo credit Aljos Farjon.

Map. SA-34. Lepidothamnus fonkii (Podocarpaceae)

[n=34, m=33, h=11, 1857–2006]

This species and Pilgerodendron uviferum

often occur together and consequently their
distribution in southernmost Chile is similar.
Lepidothamus fonkii is a creeping or more or
less erect shrub commonly up to 50 cm tall,
forming extensive vegetation mats in Sphag-
num bogs and along margins of shallow lakes
and ponds, sometimes over rocks. In Chile it
occurs from Los Lagos (Alerce Costero N.M. in
Prov. Valdivia) at 40° 11’ S to Tierra del Fuego
at ca. 55° S, making it the southernmost coni-
fer in the world. In Argentina it is known
from a few localities in Chubut and Santa
Cruz near the border with Chile. Its distri-
bution seems to be discontinuous between
44° S and 48° S but this may be in part an
artifact of the ­limited number of herbarium
collections made of this species. Similarly, it
is very likely to occur in the large archipelago
in the far south of Chile (partly included in
the Fernando de Magallanes National Park)
but no herbarium collections have come to
our attention. In the north of its range it is
growing in bogs in forested areas with Pilgerodendron uviferum, Fitzroya cupressoides, Podocar-
pus nubigenus and Nothofagus antarctica. Further south Fitzroya is absent, but the other two trees
­continue to form the fringes of bogs on higher ground. The altitude range of Lepidothamnus fonkii
is from near sea level in the far S to 930 m on Cerro Mirador in Los Lagos.
454 south america

Map SA-35. Podocarpus nubigenus (Podocarpaceae)

[n=50, m=47, h=7, 1833–2004]

Podocarpus nubigenus is distributed in Chile

from near Valdivia (Parque Oncol) at 39° 42’
S to beyond 50° S in Magallanes; in Argentina
it has been observed in Neuquén, Chubut and
Santa Cruz in localities close to the border
with Chile (not mapped). This distribution is
similar to that of Pilgerodendron uviferum and
Lepidothamnus fonkii but Podocarpus nubig-
enus does not extend as far south as these
two species. All three species reach their
northern limits in the same area, the Cordil-
lera de la Costa between Valdivia and Osorno
in Los Lagos. This coastal range has several
protected areas in which the northernmost
sections of the Valdivian coastal temperate
rainforest are preserved: Alerce Costero Natu-
ral Monument, Valdivia National Reserve and
others. The mountains here reach 700–1000
m but N of Valdivia the coastal range drops
to 300–400 m and precipitation levels drop
also. Podocarpus nubigenus is often found in
water-logged soil or near streams on acidic
substrates and can occur in forest dominated
by conifers or by angiosperms, or in a mixture
of both. A lack of collections of herbarium
specimens from Argentina in the BRAHMS
Conifer Database explains the absence of dots there. Mere observations (GBIF occurrences records,
accessed 23 April 2012) are not included in the database.

Podocarpus nubigenus on the edge of Valdiv-

ian rainforest in Parque Oncol, Cordillera de
Queule, Valdivia, Chile. This species and the
podocarp Saxegothaea conspicua are common
components of this evergreen rainforest, occur-
ring with e.g. Drimys winteri, Amomyrtus luma,
A. meli, Laureliopsis philippiana, Eucryphia
cordifolia and Lomatia dentata. The understo-
rey is dominated by ferns and the trees are
covered in epiphytes. These shade tolerant
conifers can successfully compete with the
angiosperms and reach the canopy, as this old
tree did before the forest was locally disturbed
when a road was bulldozed through it. Photo
credit Aljos Farjon.
 south america 455

Map SA-36. Saxegothaea conspicua (Podocarpaceae)

[n=62, m=61, h=10, 1830–2008]

Saxegothaea conspicua occurs in Chile from

Maule (Region VII) to Aisén (Region XI) but
is more common N of 44° S. In Argentina it is
found close to the border with Chile in Neu-
quén, Rio Negro and Chubut. This is another
conifer species common in wet Valdivian tem-
perate rainforest, most widely distributed in the
Andes but also present in the Cordillera de la
Costa especially in the province of Valdivia and
on Chiloé Island. It occurs from near sea level
in the coastal ranges to ca. 1000 m in the Andes,
where it is most abundant along streams. North
of 38° S it becomes scarce due to the transition
to a drier climate. This species is quite unique
in the family Podocarpaceae; not only is it
endemic to this region, but no fossil record from
elsewhere is as yet known. Phylogenetic analy-
sis (Biffin et al. in Turner & Cernusak, eds. 2011)
has placed it, with Microcachrys tetragona from
Tasmania, as basal to the ‘Dacrydioid’ and ‘Pod-
ocarpoid’ clades and by implication as a primi-
tive member of the family. Both monospecific
species are characterized by seed cones with multiple fertile scales which, in view of the strongly
reduced cones of all podocarps in the two subsequent clades, may represent an ancestral state.

Map SA-37. Podocarpus salignus (Podocarpaceae)

[n=69, m=65, h=8, 1833–2011]

Podocarpus salignus is widespread in the Chil­

ean regions of Maule, Biobío, La Araucanía
and Los Lagos; it does not occur in Argentina.
It occurs in the Cordillera de la Costa, the Cen-
tral Valley and the Andes. In the northern part
of its range where the climate has similarities
to that of the Mediterranean Podocarpus salig-
nus is mostly limited to water courses and cool
ravines. In the Andes it is a minor component
of Nothofagus obliqua forests or it occurs on wet
mountain slopes where it can form nearly pure
stands. The altitude range is between 20 m and
1200 m. In Los Lagos the southernmost locality
recorded from a herbarium collection at MO is
from Achao Island on the E side of Chiloé Island
at 42° 28’ S and was collected in 1982. This spe-
cies becomes more common N of Osorno at 40°
35’ S and its northernmost occurrence is near
Linares at 35° 50’ N so apart from a few southern
outliers its extent of occurrence (EOO) is limited.
Podocarpus salignus is in decline throughout its
range but particularly in the northern part due
456 south america

to habitat degradation, forest clearance and spread of plantation forestry using exotic trees. This
species is listed as Vulnerable (VU) on the IUCN Red List.

Podocarpus salignus in a forest remnant near Punucapa, Valdivia, Chile. Photo credit Aljos Farjon.

Map SA-38. Prumnopitys andina (Podocarpaceae)

[n=34, m=30, h=7, 1828–2011]

Prumnopitys andina is the southernmost spe-

cies of the genus in South America, three other
species occur in the Andes N of 30° S (see map
SA-5 on p. 432). This species has a much more
limited distribution, occurring in three regions
of Chile: Maule, Biobío and La Araucanía and
in Neuquén in Argentina near the border with
Chile. It extends from ca, 35° 50’ S to ca. 39°
30’ S and thus has a similar northern limit as
Podocarpus salignus (map SA-37 on p. 455) and
Saxegothaea conspicua (map SA-36 on p. 455);
undoubtedly determined by climate factors
and especially summer drought. Its altitudinal
range is from 200 to 1400 m and in the Andes it
mostly follows river valleys and is often accom-
panied by Austrocedrus chilensis. Prumnopitys
andina descends into the foothills of the Andes
and occurs in a few hills and ravines in the Cen-
tral Valley, but rarely as far W as the eastern
side of the Cordillera de la Costa (W of Angol in
 south america 457

La Araucanía). It is estimated that fewer than 10–12 subpopulations remain of this species, many
not counting more than 100 mature trees. Since the map is based on herbarium collections going
back to the nineteenth century, some dots may represent subpopulations that have been wiped
out by deforestation, conversion to plantation forestry or general habitat degradation preventing
successful regeneration. This species is listed as Vulnerable (VU) on the IUCN Red List.
Africa

Geography

The continent of Africa is with ca. 30.4 million km² the second largest land mass. Madagascar
adds an additional 0.6 million km². From its most northerly point, Ras Ben Sakka in Tunisia at
37° 21’ N to its most southern point, Cape Agulhas in South Africa at 34° 51’ S is a distance of ca.
8000 km and the greatest E-W distance is ca. 7400 km. Unlike in other large continents, there
are no great mountain ranges with continuous high altitude; most of the only 10 high mountains
over 4000 m are isolated volcanos. The highest mountain on the continent is Mt. Kilimanjaro, a
volcano at 5895 m. In the NW the continent is bordered by the Atlas Mountains which receive
rain from the Atlantic but also have a rain shadow side next to the Sahara. This desert, the largest
in the world with ca. 9 million km², extends from the Atlantic Ocean to the Red Sea and from
the Mediterranean Sea to the Sahel zone, a broad transition from desert to savanna and dry for-
est. Further extensive deserts are situated in southern Africa, with the Namib Desert along the
Atlantic coast and the Kalahari Desert in the interior. The most extensive uplands are the Ethio-
pian Highlands in the E, loosely connecting with mountains in Kenya and Tanzania; W of these
are some high mountains bordering the Rift Valley, such as Rwenzori (5109 m). This valley is a
long system of tectonic rift (an incipient ocean) along which large elongated lakes have formed.
Intermittent mountainous terrain extends from the Rift Valley S to the South African Cape (Table
Mountain). Most of the W half of the continent is lowland, with Mt. Cameroon (4095 m) the only
notable exception. The Congo Basin is a very large, level basin forming the drainage of the largest
river by volume in Africa, the Congo. After the Amazon Basin in South America, it is the largest
area of tropical lowland rainforest in the world. The continent can be divided into three major
geographical regions relevant to conifers: The Atlas Mountains and the Mediterranean coast, the
Afromontane region from Ethiopia to South Africa, and Madagascar. A smaller area is formed by
Mt. Cameroon and the surrounding highlands; it is the most isolated in terms of distance to other
areas with conifers. The most substantial biogeographical barrier separating the continent from
Eurasia is the Sahara; this desert also separates the Atlas Mountains and the Mediterranean coast
from sub-Saharan Africa. Within sub-Saharan Africa, the major barriers to conifer distribution
are the Kalahari Desert and the Congo Basin. The most important conduit for conifer distribu-
tion is the archipelago-like arrangement of higher altitude areas and mountains that constitutes
the eastern African Afromontane region. Towards the southern tip of Africa, a temperate climate
allows species from the Afromontane forest to descend to near sea level, but near the Cape and
especially up the W coast increasing aridity turns the vegetation from treeless fynbos to desert.
 africa 459

Conifers in Africa

Families Genera Species Species + infra Countries with Area of occupancy Endemic taxa
conifers in km²
4/8 10/70 34/615 41/794 25/54 14,625/31,000,000 22
Numbers after ‘/’ are global or continent totals.

Compared to other continents, conifers are poorly represented in Africa. Vast areas of the con-
tinent are devoid of conifers. The main distribution of conifer species is in the Atlas Mountains
and throughout the eastern African Afromontane region, with smaller areas with conifers around
Mount Cameroon and in Madagascar. The southern coast of the Mediterranean has a few spe-
cies in the less dry areas. Elsewhere, conifers are sparsely distributed across the region between
the Kalahari Desert and the Congo Basin. In the Sahara, there is only is a single population of
Cupressus dupreziana. There are no conifers in the Congo Basin, the Kalahari Desert, the highland
areas west of Mount Cameroon, the Horn of Africa south of the coastal Cal Madow range and
in the entire western parts of Madagascar. Conifers are also absent from savanna and woodland
belts in Southern Africa and south of the Sahara; and from the Karoo and other plateaus in South
Africa.
The two principal influences determining conifer distribution in Africa relate to ecology and
history. Many native conifer species in Africa have limited seed dispersal capability. Few spe-
cies have effectively winged seeds and these only occur in the Atlas Mountains (Abies, Cedrus
and Pinus species). Some species have virtually no dispersal mechanism. The Podocarpaceae in
Africa and the sole Juniperus species ( J. procera) are bird dispersed. Of these, only J. procera and
Podocarpus milanjianus have spread widely and have maintained these broader distributions. The
extreme conditions in the deserts and semi-deserts of the Kalahari, Namib, Sahara and the Horn
of Africa explain the absence of conifers in these regions. The unusual and solitary occurrence in
the Sahara of Cupressus dupreziana is a classic example of a relict population (see page 159).
Explaining the absence from the Congo Basin is more difficult. Conifers, mainly Podocarpaceae
and Agathis, do occur in lowland tropical rainforests, for example in Borneo. There are no obvi-
ous geographical barriers for conifer distribution into this basin from the surrounding regions,
especially from the east. This suggests that ecological conditions have been and are the limiting
factor. In lowland tropical rainforest environments, conifers are restricted to nutrient deficient
soils, for example, white sands (e.g. the kerangas of Borneo) and ultramafic soils. The sedimentary
nature of the Congo Basin with numerous flooding rivers heightens soil fertility throughout the
region. The growing season is virtually year-round and conditions are without edaphic limita-
tions. Under these conditions, competition with angiosperms is severe, limiting opportunities for
establishment.
These limiting conditions however do not apply in the vast stretch of tropical highlands west of
Mount Cameroon. Here, suitable habitat appears to be common yet is devoid of naturally occur-
ring conifers. This lack of conifer species must have an historical explanation. Fossil evidence
indicates a presence of conifers until the end of the Pliocene (Morley in Turner & Cernusak,
eds. 2011); presumably they went extinct during the Pleistocene. Introduced conifers from similar
habitats in Africa (e.g. Afrocarpus mannii) grow well in the region. Further underlining the pecu-
liarity of this absence is that the genus Afrocarpus, distributed widely in the Afromontane region,
is only represented by A. mannii on the island of São Tomé off the West African coast. Podocar-
pus in West Africa is only represented by the most widely distributed species P. milanjianus but
even this species does not occur west of Mount Cameroon. Extinction cannot be excluded as an
explanation as there is fossil evidence of podocarpaceous pollen in West Africa beyond the pres-
ent occurrence of members of the family Podocarpaceae.
460 africa

The greater diversity and presence of conifers in the Afromontane region can be explained by
the corridor function of this chain of loosely connected highlands and mountains. Bird dispersal
has facilitated distribution across gaps of unsuitable habitat (e.g. the Serengeti Plain) and the
lowering of Afromontane vegetation belts during the Pleistocene glacial maxima made the for-
ests more joined. Madagascar with its five species of Podocarpus is an example of relict distribu-
tions, probably related to past climate change and recent human interference. These species, all
endemic, could have evolved from a single immigrant ancestor from continental Africa.

Map AF-1. Distribution of conifer species in Africa.

 africa 461

Map AF-2. Map of Africa showing numbers of coni-

fer genera per one degree cell.

Africa has a total of only 10 conifer genera. The

cell with the largest concentration of genera
on the north coast includes Cedrus, Juniperus,
Pinus, Taxus and Tetraclinis. This area belongs to
the Mediterranean region which is not African
in floristic terms. These species are mapped and
discussed in the chapter Europe and the Medi-
terranean. In sub-Saharan Africa and Madagas-
car no cell occurs with more than three genera,
� indicating the poverty of conifer diversity in the
�
�
continent.
�
�

Map AF-3. Map of Africa showing numbers of coni-

fer species and/or taxa of lower rank per one degree
cell.

The areas with the highest number of different

species (7 species per degree cell) are on the
north coast in Morocco and Algeria. No cell has
more than seven species. In Madagascar there
is one cell with 6 taxa. The picture is essentially
the same as with the generic diversity, and again
the Mediterranean region has most of the lim-
ited diversity of conifers in Africa. The causes of
�‒� this poverty are likely to be historical, i.e. a his-
�
�‒�
tory of extinction and lack of immigration from
� Eurasia. There is not a paucity of suitable habi-
�‒�
tats, especially in the Afromontane region, that
could explain this lack of species ecologically.

Map AF-4. Rarity values per one degree cell. Cell

score is the total of the rarity calculations for each
species in the cell. The rarity calculation is 1 divided
by the total of cells occupied by a species. Thus, a
species that occurs in one cell scores 1 while a species
that occurs in 10 cells scores only 0.1.

The two highest scoring cells are on Madagas-

car. Global rarity is the primary criterion, and
this together with the number of such limited
taxa within a cell gives it a high value. Here the
Mediterranean region scores much lower than
in the previous two maps on diversity, as most of
the species there occur also elsewhere and even
outside Africa. The higher the score, the more
narrow endemism occurs in a given degree cell;
here orange cells have one taxon limited to that
cell and red cells have more than one that do
not occur outside it.
462 africa

Juniperus

Map AF-5. Juniperus procera [n=132, m=109,

h=14, 1838–2011]

Juniperus procera is of major biogeograph-

ical interest because it is the only conifer
in Sub-Saharan Africa with a close taxo-
nomic affinity to species in Eurasia. The
genus is widespread in the Northern Hemi-
sphere both in Eurasia and North Amer-
ica, but only J. procera in Africa crosses
south of the equator, ultimately to 18° S
in Zimbabwe. Its nearest sister species
is J. excelsa (Adams, 2011, p. 10), which is
distributed in the eastern Mediterranean–
Black Sea area, disjunct from the range of
J. procera due to desertification of much
of the Middle East. Connecting suitable
habitat between the two ranges probably
existed prior to climate change, enabling
migration of junipers far into Africa. Once
separated, the two taxa diverged into dis-
tinct species.
The distribution of J. procera in
Africa is itself very scattered, with iso-
lated populations in Marungu (Congo
Democratic Republic), the Nyika Plateau (Malawi) and the single ancient, most southerly
tree in Zimbabwe, possibly a relic of a larger population. These may have been established
by long distance seed dispersal carried by birds. Juniperus procera grows in the Afromon-
tane forests where it is often a dominant tree, or forms the canopy of mixed forest with other
conifers (Afrocarpus, Podocarpus) and angiosperms like Olea spp., Faurea saligna, Dombeya
mastersii and Ilex mitis. In the northern sections of the range of J. procera the species follows
the high escarpments above the deserts of Somalia and the Arabian Peninsula, bordering the
Gulf of Aden and the southern part of the Red Sea. Here there are no closed tropical mon-
tane forests, but scattered woodlands in which the juniper is a major component tree species.

Juniperus procera woodland near Yavello in S Ethiopia. Photo credit Ib Friis.

 africa 463

Widdringtonia

Widdringtonia belongs taxonomically to a group of genera (Callitrideae) in the family Cupres-

saceae that is distributed in the Southern Hemisphere. Their closest relatives are possibly Actinos-
trobus and Callitris in Australia (Farjon, 2005a). Widdringtonia is one of only two genera endemic
to Africa (the other is Afrocarpus, see map AF-11 on p. 465). However, there is a fossil record from
North America, dating to the Late Cretaceous (Cenomanian, 95 Ma) that has been described
as a species in this genus (McIver, 2001). This is an intriguing discovery, as it appears to dem-
onstrate that Widdringtonia was not restricted to the southern hemisphere in the past. All its
recent closest relatives are in the southern hemisphere, but these may have had links with taxa
in the Northern Hemisphere, too. Extinction has made the reconstruction of past distributions
very difficult and confirmation by well identified fossils is very rare. A few fossils from Tertiary
lignite beds indicate that the genus occurred in locations in South Africa from where it has since
disappeared (Barker & al., 2004).

Map AF-6. The distribution of the

genus Widdringtonia.

The genus is endemic to Africa

and occupies the southern part
of the Afromontane region. It
has four species, W. nodiflora, W.
cedarbergensis, W. schwarzii and
W. whytei. Only W. nodiflora is
widespread and occupies nearly
the entire range of the genus,
occurring in Malawi, Mozam-
bique, Zimbabwe and South
Africa. The other three species
are highly localized. Two are
endemic to the Cape Province of
South Africa (W. cedarbergensis and W. schwarzii) and the third species, W. whytei, is endemic
to Mount Mulanje in Malawi. The three localized species are all in danger of extinction (IUCN
categories CR, VU and EN respectively) due to recent and continuing decline. Widdringtonia nodi-
flora is not threatened and considered to be Least Concern (LC).

Map AF-7. Widdringtonia nodiflora [n= 116, m=89, h=13, Map AF-8. Widdringtonia cedarbergensis [n=
1738–2005] 20, m=11. h=20, 1839–1982]
464 africa

Map AF-9. Widdringtonia schwarzii [n=15, m=14, h=10,

1904–2005]

Map AF-10. Widdringtonia whytei n=14,

m=14, h=7, 1891–1998]

Three of the species are extremely limited but one is widespread. What could be the cause of
this? The best explanation is probably found in the different ecology (Pauw & Linder, 1997). The
three restricted species are trees with limited adaptations to fire. These have a similar strategy
to species of Cupressus (see pages 76–78) and of many pines (Pinus spp.). Trees are often killed,
but the serotinous cones protect the seeds if the fire is not too hot and these are afterwards dis-
persed and germinate with the coming of winter rains. But if fires become more frequent, seed-
lings and saplings are killed too before they become mature. Drying climate and increasing fires
must have caused these species to retreat to high mountain summits or deep, protected gorges,
which is indeed where they are found now. Widdringtonia nodiflora has an additional adaptation
that is absent in the other species: it coppices after fire. Trees sprout with multiple stems from
an undamaged root base. In fire-prone environments, many trees, e.g. Australian eucalypts, can
regenerate in this way. As a result, W. nodiflora could, unlike the other species, maintain much of
its wide distribution across southern Africa.

Widdringtonia cedarbergensis was once abundant
in the Cedarberg Mountains of the Western Cape.
Logging replaced the forest with the flammable
fynbos vegetation and now the only remaining
trees occur among the bare rocks where fires do
not reach. It is Critically Endangered (CR) on the
IUCN Red List. Photo credit Aljos Farjon.
Widdringtonia whytei occurs on Mount Mulanje in
Malawi in isolated groves and in steep gorges where it
is more or less safe from fire. People increase the fires to
stimulate grassland; they also (now illegally) cut trees
for the valuable timber. On this mountain, W. nodiflora
also occurs and could under these conditions eventu-
ally replace W. whytei. Photo credit Philip Cribb.
 africa 465

Afrocarpus

Map AF-11. The distribution of the genus Afrocarpus.

Afrocarpus (Podocarpaceae) is a small genus of five species endemic to the African continent.
It is closely related to Podocarpus, which has a pantropical distribution. Some botanists have
included Afrocarpus in that genus but there are morphological and molecular differences (Barker
et al., 2004) supporting its separate status at this rank. All species potentially grow to large trees
and are sometimes co-dominant and in other forest types form part of a more mixed canopy,
where they are often emergents. They are late-successional and long living trees and the seeds
form ‘imitation fruits’ eaten and dispersed by birds. The five species are: A. dawei, A. mannii, A.
usambarensis, A. falcatus and A. gracilior. Of these, the best known is the South African A. falcatus
or Bastard yellowwood.
The distribution of the genus Afrocarpus appears to be divided into a northern and a southern
area separated by a gap of 1400 km (1800 km if we ignore the solitary occurrence in northern
Malawi). Far to the west, on the island of São Tomé (red circle), A. mannii is the only natural rep-
resentative of the genus in West Africa. In East Africa, the genus extends north into the Ethiopian
Highlands (where Podocarpus is absent) and is widespread in the higher parts of Kenya, Tanzania,
and in the highlands or swamps west and northwest of Lake Victoria (where it is often joined by
Podocarpus). Beyond the gap to the south it reappears in South Africa (including the southern
tip of Mozambique) where it extends all the way to the Cape. At the Cape, these trees descend
to near sea level, but further north and in all of East Africa they are limited to highland habitat.
This imposes limitations to the distribution and explains some of the patterns, but not the great
gap. There are highlands in southern Tanzania, Malawi, Mozambique and Zimbabwe where Podo-
carpus occurs (see map AF-17 on p. 468) but Afrocarpus is absent. The habitat in Africa of these
two genera is mostly similar and they often occur together in the forest. Fragmentation of the
Afromontane forest, which was once much more continuous (Huxley et al. eds. 1998), may have
466 africa

reduced ranges and area of occupancy but would not explain the gap. The gap is weakly bridged
by the occurrence of A. falcatus, the South African species, in Northern Malawi (see map AF-14
on p. 467), which forms a far outlier of that species. A phylogeny of Afrocarpus might throw some
light on the issue by indicating lines of descent and possibly where the genus originated. This has
not been researched to date. The absence of Afrocarpus in Madagascar has been confirmed whilst
researching the herbarium data for this Atlas: records there refer to planted trees only and it has
not been seen in the wild.
The occurrence on São Tomé appears to demonstrate that seed dispersal by birds must play a
major role in the distribution patterns of this genus. This island in the Gulf of Guinea is of vol-
canic origin and only ca. 1 million years old; the establishment of a forest on its highest summit,
where this species occurs, is much more recent. Either this endemic species derived from a now
extinct ancestor in the region, e.g. in the Mt. Cameroon area, or from a species in East Africa;
in the latter scenario dispersal across the Congo Basin to the west coast is involved unless it is
assumed that in a cooler climate podocarps could have existed there. The single occurrence of
A. falcatus in Malawi could also be the result of dispersal. It would appear then that the patterns
of distribution seen today are primarily historical, involving random dispersal events, followed by
spread where ecologically possible from such centres of origin.

Map AF-12. Afrocarpus gracilior [n=125, m=104, h=4, 1863–2007]

Afrocarpus gracilior is the most widespread species

in the genus. It is common in the mixed conifer-
angiosperm forests of the East African Highlands
including Ethiopia, where it becomes the only podo-
carp conifer. Its altitude ranges from 1500m to 2600 m
a.s.l. Some forests on the highest ridges are dominated
by this species and Juniperus procera and these are the
closest to a natural conifer forest we can observe in
Africa, south of the Sahara. The areas on three sides
of Lake Victoria occupied by savanna or swamp for-
est are unsuitable for this species, hence the large gap
between the main distribution and the western popu-
lations around Lake Kivu.
Afrocarpus gracilior in Ethiopia. In the
tropics, solitary trees like this one are
usually not planted, they are left-overs
from the forest which has been cut and
the land converted to pasture, possibly
less than a century ago. Cattle need
shade trees in the tropical sun, and
trees like this were left for that purpose.
Such a tree may be the source of the
herbarium collection on which a dot on
the map is based. Therefore, one cannot
extrapolate from the map the present
extent of the forest of which this species
is a common component. Photo credit
John Grimshaw.
 africa 467

Map AF-13. Afrocarpus usambarensis [n=51, m=41,

h=4, 1893–2005]

Afrocarpus usambarensis occurs in the Usam-

bara and Paie Mountains of Tanzania, the
Chyulu and Taita Hills of Kenya, and in isolated
highland areas elsewhere in Tanzania. Exten-
sive grassland and savanna areas such as the
Masai Steppe and the Serengeti Plain separate
these populations. The forests are evergreen,
dry or wet, and mostly composed of various
angiosperms. Trees of this species are often
solitary but not emergent, only reaching into
the general canopy of the forest. In wet forest it
is joined by another conifer, Podocarpus milan-
jianus. The drier forest type, at lower altitudes, is often degraded and converts to coarse grassland
under human influence. This exacerbates the island character of the distribution of this species.
Logging in the wet rainforest of the two conifers reduces the slower growing Afrocarpus to a rare
tree in remaining forest patches. It is classified as Vulnerable (VU) on the IUCN Red List.

Map AF-14. Afrocarpus falcatus [n=41, m=39, h=4,

1775–2005] Map AF-15. Afrocarpus dawei [n=33, m=27, h=3,
1907–1997]
Afrocarpus falcatus occurs along the south
Afrocarpus dawei is found in northern Tanzania
coast of South Africa and is scattered in the
and in Uganda in seasonal swamp forests. This
highlands further north, just in Mozambique
flatland region (1100–1200 m a.s.l.) is inundated
in coastal locations; an outlier population is
by slow running rivers that drain into Lake Vic-
known from Northern Malawi on the Nyika
toria. In the rainy season these forests can be
Plateau, where it is joined by another dis-
extensively flooded and inaccessible. Interest-
junct Afromontane conifer, Juniperus procera
ingly, this region is outside the Afromontane
(map AF-5 on p. 462). Limited to moist valleys
forests and has a distinct flora. This species is a
with permanent rivers, or sheltered ravines, it
co-dominant most commonly associated with
becomes both scarcer and a smaller tree going
Baikiaea minor (Leguminosae) and Mimusops
northward. In the coastal Knysna Forest of the
sp. (Sapotaceae), or a more solitary emergent.
Cape some giant trees maybe 1000–1500 years
The scattered dots on map AF-15 are more an
old and here this species is a large emergent
indication of the inaccessibility of the swamp
in the forests bordering the rivers, perhaps a
forests than of the rarity of this tree. Half of all
reminder of a wetter climate in South Africa
herbarium collections (16) in the Conifer Data-
millions of years ago. The Afromontane forest
base used to produce the map came from the
descends here to the ocean coast.
Inziro Forest Reserve in Tanzania, to which
there is a road.
468 africa

Map AF-16. Afrocarpus mannii [n=3, m=2,

h=2, 1861–1996]
Afrocarpus mannii is the most narrowly
distributed species in the genus and one
of the very rare conifers of Africa (see
map AF-4 on p. 461). Its sole locality, the
summit area of Pico, on the shield vol-
cano that forms the island of São Tomé,
is ca. 2300 km west of the nearest loca-
tion of another species in the genus. It
occurs there from ca. 1450 m to the sum-
mit area at 2142 m a.s.l. It is nowhere a
tall tree and at the summit it is reduced
to dwarfed krummholz. It is common
in the high montane cloud forest where
this has remained undisturbed. This spe-
cies is listed as Vulnerable (VU) on the
IUCN Red List. Deforestation at lower to middle altitudes on the mountain is the main threat to
this species.

Podocarpus

Map AF-17. The distribution of

the genus Podocarpus in Africa.

The pantropical to Southern

Hemisphere genus Podocarpus
(Podocarpaceae) has a surpris-
ingly limited presence in Africa
and Madagascar with four spe-
cies on the continent (P. milan-
jianus, P. latifolius, P. henkelii
and P. elongatus) and another
five endemic to Madagascar
(P. capuronii, P. humbertii,
P. rostratus, P. perrieri and
P. mada­gascariensis, the latter
with three recognised variet-
ies). On the continent, aside
from the more widespread
Podocarpus milanjianus, these
are broadly confined to the Afromontane spine in the East and South. The species of Madagascar
are endemic, narrowly restricted and generally endangered, especially P. perrieri which is only
known from the type and one other location. Deforestation and habitat loss threatens most if not
all on Madagascar. The limited distribution and spread of Podocarpus in Africa is discussed in the
introductory section of this chapter.
 africa 469

Map AF-18. Podocarpus milanjianus

[n=284, m=207, h=10, 1891–2007]

Podocarpus milanjianus is by far

the most widespread species in
Africa of this genus; it is among
the few species with a near conti-
nental range in the family Podo-
carpaceae. The main range of this
tree is in the East African high-
lands of Kenya, Tanzania and in
the mountains surrounding the
Western Rift Valley. Peculiarly,
the genus, and thus also this spe-
cies, is absent from the highlands
of Ethiopia; its northernmost
population in East Africa is in the
Imatong and Dongotona Moun-
tains in the far south of Sudan. Podocarpus milanjianus is a montane to high montane species
occurring in tropical evergreen rainforest, cloud forest, or at its highest limit in dwarf forest domi-
nated by Ericaceae, interfacing with subalpine grassland. It does not form conifer forest, although
it can occur in forest dominated by Juniperus procera in e.g. Kenya, and is usually accompanied
by angiosperm trees. The distribution is therefore naturally limited to areas with high mountains,
which receive high levels of rainfall in a tropical climate. It ranges from 8° 24’ N to 19° 59’ S and
is therefore confined to the tropics, although it occurs at high altitude, usually between 1300
and 3000 m. A more scattered distribution extends the range of P. milanjianus to western Africa,
where another concentration is found in the Cameroon Highlands. The link between East and
West Africa for this species follows more or less isolated highland areas to the south of this basin
in Zambia and Angola, but as far as present knowledge based on herbarium collections goes,
apparently with wide intervals. A dispersal route along these lines was suggested by Maley et al.
(see map reproduced in Turner & Cernusak, eds. 2011, p. 35).

Map AF-19. Podocarpus elongatus (red) [n=19, m=18, h=3, 1815– Podocarpus latifolius at Laager Farm,
1970]; Podocarpus henkelii (black) [n=30, m=27, h=6, 1916–1988]; KwaZulu-Natal, South Africa, at 750
Podocarpus latifolius (yellow) [n=38, m=34, h=4, 1815–1989] m a.s.l. in half-open woodland. Photo
credit G. R. Nichols.
470 africa

Podocarpus elongatus occurs in the Eastern, Northern and Western Cape Provinces of South
Africa, Malawi, Zambia and Zimbabwe growing at elevations between 130 and 2250 m. It is an
uncommon species growing in woodlands in moist sites, usually along intermittent streams and
in ravines, or on rocky sites with sparse vegetation. In the Western Cape its habitat abuts on
fire-prone vegetation types such as fynbos and as a consequence individual trees are frequently
burnt. Resprouting from the base, they then develop into broad spreading shrubs or bushes, while
only individuals that are protected from fire, e.g. by growing in a deep ravine, can develop into
monopodial trees of some size, up to 20 m tall. In the NE part of its scattered range it is usually
a component of moist evergreen forest and grows more often into a tree.
Podocarpus henkelii occurs in the Eastern Cape Province and Kwazulu Natal in South Africa,
Malawi, Tanzania and Zimbabwe. Podocarpus henkelii occurs in montane evergreen rainforest,
often on steep, rocky slopes, at altitudes between 1300 m and 2000 m. It is also present in coastal
forests near sea level in Eastern Cape Province (where it is rare) and Kwazulu Natal. Here it
may be associated with Afrocarpus falcatus; both are in these forests emergents above a canopy
of angiosperm trees. In Tanzania it occurs as a codominant in Ocotea-Podocarpus forest with a
canopy 30–40 m tall and both Afrocarpus usambarensis and P. milanjianus are often also present
as canopy trees. The global population of this species is extremely fragmented according to evi-
dence from herbarium collections, only occurring in five widely separated locations. One of these
is in the eastern part of Tanzania where podocarps are intensively logged. In South Africa the sub-
population is more stable but the species is reported to be rare. Information about trends in the
two known locations in Malawi and Zimbabwe is lacking. Unsustainable logging poses the major
threat to this species, especially in Tanzania. This species is listed as Endangered (EN) on the
IUCN Red List. In South Africa and Zimbabwe it is commonly planted as an ornamental tree.
Podocarpus latifolius is known from the Cape to the Northern Transvaal in South Africa. This
species is a canopy forest tree in the coastal and midland primary forests where there is sufficient
rainfall and natural protection from fires to allow such forest types to develop. In open coastal
bushland and on dry, rocky mountain slopes it only grows to a stunted tree a few meters tall at
most. Elevation range is from sea level up to ca. 2000 m. The tree was extensively used in colonial
times for railway sleepers and construction, and many houses were built with it, such as the old
Cape homesteads of which many still exist. In South Africa this species is commonly planted as
an amenity tree in parks and along streets. Elsewhere it is uncommon and mainly represented by
specimens in botanic gardens. It is too slow growing for profitable forestry plantation.

Podocarpus in Madagascar

Map AF-20. Podocarpus capuronii [n=17, m=17, h=4, 1919–2004]

Podocarpus capuronii is probably now restricted to the Itremo

Massif, where all recent collections have been made. The dis-
junct occurrences are based on Perrier de la Bathie 13163 in the
Paris Herbarium (P), which was collected in the 1920’s (exact
date unknown) and on R. Capuron 27065, the type of P. woltzii
(P) in the N of Madagascar, also undated. Deforestation, habitat
loss, cutting for firewood, fires, and grazing are the main threats
to this species. This species is listed as Endangered (EN) on the
IUCN Red List. Podocarpus capuronii is found on skeletal sandy
soil over quarzite or gneiss, along streams in ravines and on rocky
slopes and ridges. It is a slow growing shrub or stunted tree in
these depauperate sites; in forest it may attain 20 m. Altitude
ranges from 1320–2000 m, but is reported to extend to 2800 m.
 africa 471

Map AF-21. Podocarpus humbertii [n=10, m=10, h=4, 1924–2001]

Podocarpus humbertii is known from sub-humid forest, dry

lowland deciduous forest, and ericoid thickets or wooded
heath on mountain summits of gneiss and granite in Antsira-
nana Province with elevation ranging from ca 1600 m to 2410 m.
In the original description (De Laubenfels, 1972) it was said to
occur up to 2800 m. Especially the sub-populations in lowland
forest have been affected by deforestation and this species is
listed as Endangered (EN) on the IUCN Red List.

Map AF-22. Podocarpus rostratus, [n=10, m=9, h=4, 1912–2005]

Podocarpus rostratus, recorded in the past from Antsiranana,

Fianarantsoa, Mahajanga and Toamasina Provinces, is now
known from only two highly disjunct populations and is clas-
sified as Endangered (EN) on the IUCN Red List. It occurs in
ericaceous scrubland near the summits of mountains on rocky
slopes on silicious rocks within elevation range 1800–2400 m.
472 africa

Map AF-23. Podocarpus perrieri, [n=3, m=3, h=4, 1924–1951]

Podocarpus perrieri, known to have occurred in Fianarantsoa

and Antanarivo provinces, is now only known from its type
location in the Forêt d’Andasibé (Toamasina, Antanarivo). It
was collected by H. Humbert in the Andringitra Massif in 1924
but has not been found there again, while a third collection
made in 1951 not far from the type location is of uncertain iden-
tification. This species occurs in tall, subhumid forest and is
reported to have attained 30 m. Altitude ranges likely between
1200 and 2000 m. This is a species on the brink of extinction,
and therefore classified as Critically Endangered (CR) on the
IUCN Red List.

Podocarpus madagascariensis has three varieties in Madagascar. The species is mostly a tree
of wet tropical montane forest, but can be found in (remnants of ) lowland forest as well as in
stunted high montane forest on mountain ridges, from level plains near sea level to altitudes of
2000–2400 m a.s.l. In lowland forest the most common variety attains tree size to 25 m tall.

Map AF-24. Podocarpus mada­ Map AF-25. Podocarpus mada­ Map AF-26. Podocarpus mada-

gascariensis var. madagas- gascariensis var. rotundus [n=3, gascariens var. procerus [n=8,
cariensis, [n=29, m=23, h=4, m=2, h=2, 1923–1996] m=8, h=4, 1934–2000]
1880–2009]

Podocarpus madagascariensis var. madagascariensis the most widespread variety, is found along
the eastern plateaux and in mountainous areas. It is a forest tree and was once abundant in
lowland rain forest; it is now more common at higher altitude where it remains a small tree.
Collections of P. madagascariensis var. rotundus are known with occurrences around Massif de
 africa 473

Bekolosy and Massif du Manongarivo. Its taxonomic distinction is somewhat doubtful. Podocar-
pus madagascariens var. procerus was listed as Endangered in a pre-1994 IUCN Categories and
Criteria listing on the grounds of its rarity and restricted distribution near Tolanaro (Fort Dau-
phin) in the SE of Madagascar. However, it is also known from the Massif de Bekolosy in the far
north of the island, and a locality in Fianarantsa Province. Its altitude ranges from near sea level
to ca. 1000 m. It has recently been reassessed but remains Endangered (EN) on the IUCN Red List
due to adverse human pressures.
 Examples of further research

The information about conifer distribution presented in this Atlas and the Conifer Database
linked to it (http://herbaria.plants.ox.ac.uk/bol/conifers) can contribute to further research in the
fields of biodiversity, biogeography, phylogeography, evolution and conservation. In this chapter
we present some examples, including two examples of research by others using the same data.
All are examples of research beyond what has been presented in the preceding chapters of this
Atlas. There are many other ways of analysing the data, depending on the questions researchers
may have. The following examples are presented in this chapter:

Analysis of the collection dates

Comparison of continents
Patterns of distribution
Conservation
GIS analysis: deforestation in Sumatera
Diversity modelling
Species distribution modelling in a phylogenetic context
The Conifer Database online

Analysis of the Collection Dates

The collection records in the Conifer Database can be analyzed by collection dates. Here we pres-
ent the 5349 dated herbarium collections of the family Podocarpaceae ordered by decade. The
most prolific decade is 1961–1970 with 1108 collections. The numbers increase from 1921 to 1940,
with a dip due to World War II; in 1951–1960 there is a steep rise to the peak in the next decade,
in 1971–1980 the numbers fall back to the level of 1951–1960. Thereafter a steep and steady decline
continues until the second decade of the 21st century. Since 1970 the number of collections made

total
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Collections in the Conifer Database of Podocarpaceae, ordered by decade. Note: the data do not extend
beyond May 2012 while many herbarium collections made during 2011 would not have been processed
when entries to the database for the Atlas project ceased. Undated collections are omitted.
 examples of further research 475

Map EFR-1. Distribution of dated herbarium collections of Podocarpaceae made prior to 1980 (blue) and
from 1980 and later (red).

per decade has fallen from 1100 to 400. In the three decades 1951–1980 a total of 2847 collections
was made, or 40.8% of the total. Since 1981 1664 collections were made, 23.8% of the total. This
means that 76.2% of all dated collections of Podocarpaceae in the database are older than 30 years
(from 2011).
How has this collecting of Podocarpaceae been distributed geographically? In map EFR-1 we
have separated collections prior to 1980 from those made in 1980 and later.
Herbarium collections from 1980 and later are well represented in Central America, the Andes
N of 30° S, Madagascar and Viet Nam. They are less abundant in S Chile, Brazil, China and Aus-
tralia and scarce in Africa, Malesia, Japan, the tropical SW Pacific and New Zealand. The decline
in collecting effort is not evenly distributed and is particularly evident in Africa and Malesia.
There is a potential significance for the accuracy of the distribution maps in those areas where
species of Podocarpaceae are declining as a result of deforestation. If a majority of map points
refer to herbarium collections made prior to 1980, a proportion of these could depict historical,
not actual presence. This possibility is further investigated in a small case study on Sumatera
below.

Comparison of Continents

Each chapter on ‘continents’ in this Atlas starts the description of the general distribution and
diversity of conifers with some statistics. Here we present these together in Table 6, in the
sequence of the chapters, for comparison.
Comparison of the values calculated for the eight ‘continents’ is not straightforward because the
areas (continents, countries within) are not equal in size. The number of countries with conifers
versus the number without is skewed as a result of this inequality. The area of occupancy (AOO)
calculated as the total area of all map points where each map point equals 5 km² could be a more
standardized measure. However, this figure rises with the number of taxa mapped and is also
influenced by the sampling density, which can vary with the availability of herbarium records. If
we compare North America and South America, roughly equal in area (as here defined) we can
see this: 126 taxa in North America give an AOO of 106,850 km² and 35 taxa in South America
give an AOO of 15,100 km². A third factor that influences the AOO is the range of a species (EOO);
North America has 13 species with very wide ranges, South America has none. It requires more
map points to cover the distribution of a wide ranging species. Despite these caveats, some com-
parisons can be validly made if we correct for the biases. If we divide the total land area by the
number of species we find Mexico, Central America and the West Indies to be the most diverse
‘continent’ with 1 species/27,800 km², followed by Malesia with 1/37,800 km² and Australasia with
1/76,600 km². Mainland Asia and Japan with the highest number (202) of species scores a mere
476 examples of further research

1/230,900 km² and North America with just 99 species is more diverse with 1/194,600 km². Europe
and the Mediterranean is much less diverse than these with 1/367,800 km² but the really poor
continents are South America with 1/525,000 km² and Africa with 1/911,800 km². These figures
suggest that the highest diversity of conifer species at this map scale occurs in regions with archi-
pelagos and small land masses at low latitudes. Some limited areas such as the California Floristic
Province in North America and the Sichuan/Yunnan mountains in China boost the diversity of
large continents, but overall these are much less diverse. Archipelagos stimulate speciation by
isolation as genetic exchange decreases with increasing distance between islands (Mac Arthur &
Wilson, 1967). Isolated mountain ranges can be ecological islands, but most mountain ranges on
the continents are connected to one another.
Endemism in areas of continental size is naturally high. Africa is lowest with 53.7%; the other
continents range between 70.5% for Europe and the Mediterranean to 91.4% for South America
and 91.0% for Australasia, which are the most isolated continents. However, North America and
Mainland Asia and Japan follow closely with 89.7% and 90.5% respectively. Africa scores low
because many species N of the Sahara are shared with Europe and the Mediterranean. With the
exception of one, all species of sub-Saharan Africa are endemic. The endemism level for North
America is remarkable given the long border with Mexico, here treated separately from it. The
same applies to Mainland Asia and Japan which has a long border with Europe and the Mediter-
ranean. It seems that at this scale the isolation versus connectivity of continents may be of little
consequence to the level of endemism and that the concept is of limited value at this scale.

Table 6. Statistics of ‘continents’ as defined in the Atlas compared.

North America

Families Genera Species Species + infra Countries with Area of occupancy Endemic taxa
conifers in km²
3/8 17/70 99/615 126/794 2/2 106,850/19,269,000 113

Mexico, Central America and the West Indies

Families Genera Species Species + infra Countries with Area of occupancy Endemic taxa
conifers in km²
4/8 11/70 99/615 132/794 23/34 55,525/2,751,100 100

Europe and the Mediterranean

Families Genera Species Species + infra Countries with Area of occupancy Endemic taxa
conifers in km²
3/8 9/70 41/615 61/794 52/53 51,575/15,080,000 43

Mainland Asia and Japan

Families Genera Species Species + infra Countries with Area of occupancy Endemic taxa
conifers in km²
6/8 36/70 202/615 283/794 39/45 107,925/46,648,350 256

Malesia

Families Genera Species Species + infra Countries with Area of occupancy Endemic taxa
conifers in km²
6/8 13/70 80/615 85/794 7/7 26,850/3,021,630 63
 examples of further research 477

Australasia

Families Genera Species Species + infra Countries with Area of occupancy Endemic taxa
conifers in km²
5/8 26/70 119/615 122/794 7/11 42,342/9,115,000 111

South America

Families Genera Species Species + infra Countries with Area of occupancy Endemic taxa
conifers in km²
3/8 9/70 34/615 35/794 9/11 15,100/17,850,420 32

Africa

Families Genera Species Species + infra Countries with Area of occupancy Endemic taxa
conifers in km²
4/8 10/70 34/615 41/794 25/54 14,625/31,000,000 22
Numbers after ‘/’ are global or continent totals.

Patterns of Distribution

Conifer species have widely divergent ranges and frequencies of distribution, with extremes
between a locality of a few hectares containing a few individuals (e.g. Pinus squamata, map
MAJ-37 on p. 202) to circumpolar and ubiquitous (Juniperus communis, maps EM-31–32 on
pp. 160–161). In the Species Index table (pp. 501–511) distributions have been compared using
the number of degree squares that each species scores on the maps presented in the Atlas. This
is inherent in the nature of the data, not a precise figure and several species (in particular those
marked with *) are estimated too low. Despite these limitations, the table provides reliable statis-
tics on relative sizes of ranges and frequency of occurrence (EOO and AOO combined).

Table 7. Number of species in five range classes for the eight ‘continents’ distinguished in the Atlas. [N.B.
Totals are not equivalent to figures given in the statistics of ‘continents’ above (species totals) because some
species occur across boundaries but have only one range and some nothospecies are not mapped.]
Degree squares/Continents 1 2–10 11–40 41–100 >100
North America 3 18 28 26 12
Mexico, Central America & West Indies 6 31 28 7 1
Europe & Mediterranean 2 10 12 7 8
Mainland Asia & Japan 14 48 72 26 6
Malesia 8 33 23 6 3
Australasia 11 69 24 5 0
South America 1 14 18 1 0
Africa 2 8 5 2 0
Total 47 222 210 80 30

Species with very limited ranges falling entirely within one degree square occur mostly in Mainland
Asia & Japan, Australasia, Malesia, and Mexico, Central America & West Indies; they are rare in
North America, South America, Africa and Europe & Mediterranean. In some cases, the limited
ranges may be an artefact of limited collecting effort; this particularly applies to some species of
Podocarpaceae in Malesia. Most species with very small EOO and AOO are genuinely restricted
and often constitute relict populations which are all that remain of a (much) wider occurrence in
the past. If range contraction and extinction are major causes for these very limited distributions,
478 examples of further research

the low numbers in some continents are probably an indication of high rates of extinction of
conifers. The climatic history of Africa and Europe in Neogene times is in agreement with this
hypothesis; in Europe it is further corroborated by the well-studied fossil record.
At the other end of the scale are very wide distributions, here defined as exceeding 100 degree
squares. Most of these are in North America, Europe & Mediterranean and Mainland Asia &
Japan and all but three are entirely in the Northern Hemisphere and none are exclusively in the
Southern Hemisphere. The skewed division of land masses between the hemispheres may play a
role, although 100 degree squares as a lower limit is not excessively large and would fit with much
room to spare in Africa, Australia and South America. The species of the northern boreal forests
amount to 12 of the total or ca. 40% of the very widespread conifer species. Three very widespread
species occur across the tropical latitudes and beyond in Malesia, and 16 are distributed in temper-
ate latitudes of the Northern Hemisphere (the ambiguity is caused by Juniperus communis which
extends across continents and from boreal to temperate latitudes but is counted under Europe &
Mediterranean).
Australasia has a disproportionately high number of species with small ranges (2–10 degree
squares), 69 or 63% of those mapped in this region. This is partly due to the endemics on rela-
tively small islands (e.g. New Caledonia, Tasmania) but mainland Australia also has a high num-
ber of species with limited distribution. Malesia, largely an archipelago with large and small
islands, has fewer than half the number of species of Australasia in this category, 33 or 45% of
those mapped in Malesia. The Malay Archipelago of Alfred Russel Wallace, while diverse like
Australasia, has fewer narrow endemics. The main reason for this is a greater connectivity of its
islands and with areas beyond in the period since the Miocene (Hall, 2009b), enabling the spread
of conifer species.
We have calculated the distribution of taxonomic diversity across the world by latitude; here
we present an example calculated for 10 degree latitude bands for genera and taxa at lower ranks
(species, subspecies and varieties) for all conifers (map EFR-2). As is to be expected, diversity
increases from high latitudes to low latitudes, but in the Northern Hemisphere the peak for coni-
fers lies between 20° and 40° N for genera (30 genera) and between 30° and 40° N for lower taxa
(320 taxa). Towards the Equator diversity drops sharply to 14 genera and 75 lower taxa. In the
Southern Hemisphere we see a gradual rise in lower taxa to the 20°–30° S band, a moderate drop
between 10°–20° S, but a rise between the Equator and 10° S to a value (76) very similar to the
same band in the Northern Hemisphere. The greatest diversity of genera in the Southern Hemi-
sphere lies at the 30°–40° S band and also shows a slight rise nearest the Equator. The diversity
of angiosperms increases with decreasing latutides right up to the Equator in both hemispheres;
The same trend also applies to pteridophytes (Moran, 2004).

Taxa Genera
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Map EFR-2. Map of the world with 10 degree latitude bands showing the global distribution of all conifers.
The taiga forest in the boreal forest region includes only a scattering of points. The scales Taxa and Genera
indicate the total numbers of taxa (species, subspecies and varieties) and genera on a latitudinal gradient.
 examples of further research 479

The trend in conifers therefore differs from other vascular plants. The high totals of lower taxa at
moderate latitudes in the Northern Hemisphere are correlated with much available land mass,
which is probably one reason why these figures are much lower at similar latitudes in the South-
ern Hemisphere. However, in the Southern Hemisphere available land mass is highest from the
Equator to 10° S and not much lower at the same band N of the Equator. At these latitudes diver-
sity is high in Malesia but very low in South America and Africa. Here the absence of conifers in
the Amazon and Congo Basins determine the low totals. Angiosperms reach maximum diversity
for their respective continents there and if conifers once occurred there, they have been out-
competed.

Conservation

The Atlas uses the listings in the IUCN Red List (International Union for the Conservation of
Nature) (http://www.iucnredlist.org/) for all conifers, which have been recently re-assessed by the
Conifer Specialist Group of IUCN/SSC (Species Survival Commission) under the chairmanship of
the first author (AF). The IUCN category of threat for each species is given in the Species Index
(pp. 501–511). These can be summarized as follows:

Table 8. Numbers of species in the different IUCN Red List Categories.

CR EN VU NT LC DD NE Total
(Critically (Endangered) (Vulnerable) (Near (Least (Data (Not
Endangered) Threatened) Concern) Deficient) Evaluated)
28 103 80 93 296 7 8 615

The three categories of threat are CR, EN and VU and together 211 species or 34% fall under these
categories. This means that more than a third of all conifer species in the world are in various
degrees threatened with extinction. We can use the information on distribution of species in the
Atlas to analyse these figures in relation to biogeographic variables such as global distribution or
range size.
We have investigated the distribution of taxa in IUCN Red List categories by mapping them
according to their listings. Here we give an example using the political units in the World Geo-

Map EFR-3. Map of the world with TDWG Standard World Geographical Scheme for Recording Plant Dis-
tributions Level-4 units (countries, provinces and states) showing the distribution per unit of taxa listed
under IUCN Red List threatened categories (VU, EN, CR). The colours from pink to red indicate an increase
in Red List significance according to the calculation ∑ (nVU1 + nEN2 + nCR3) in each Level-4 unit, whereby
n = number of taxa and categories of threat are progressively weighted. Geographical units with neutral
colouring do not contain taxa in any threatened category.
480 examples of further research

graphical Scheme for Recording Plant Distributions of TDWG (Taxonomic Databases Working
Group http://www.nhm.ac.uk/hosted_sites/tdwg) and mapping only the three categories of threat
VU, EN and CR. These categories are weighted to emphasize areas with concentrations of taxa at
risk of extinction. By using political units it becomes apparent which countries, provinces or states
(as identified at Level-4 in the TDWG scheme) have one or more species (or taxa at lower rank) in
these IUCN categories of threat. This has political significance, but does not give a detailed map
showing where threatened species occur. The maps of these threatened species can be found in
the Atlas by using the information in the Species Index.

Map EFR-4. Map of the world with the distribution of threatened (pink and red) and non-threatened
(white) conifer taxa according to their categories of threat on the IUCN Red List in 1 degree cells. Red List
significance in the threatened categories (VU, EN, CR) is indicated in two classes, with red the highest based
on a similar calculation as in map EFR-3.

The global distribution can be presented in the same manner as the diversity analysis maps in
this Atlas by using units such as 1 degree cells. Map EFR-4 gives a more detailed distribution of
threatened taxa than map EFR-3 and highlights those areas with the greatest significance (red).
The red cells are located in California, Mexico, Hispaniola, China (including Taiwan), Japan, Viet
Nam, Borneo (Sabah) and New Caledonia. When pink cells are included this map is congruent
with map EFR-3 using the TDWG political units.
Based on Red List categorization, an analysis of our data indicates one degree cell containing
37 species of which 3 are classified as Critically Endangered (CR), 11 as Endangered (EN) and six
as Vulnerable (VU). This cell is located in the Montagne des Sources/Rivière Bleue area of SE New
Caledonia. Such data analysis can be used in the planning of protected areas.

GIS Analysis: Using the Conifer Database to Assess the Decline of Conifers in Sumatera
Caused by Deforestation

Elena Shishkova, Fiona Spooner, Nicola Stanek & Lisa Wheeler

Herbarium collection data of the conifers occurring in Sumatera were extracted from the Conifers
of the World website http://herbaria.plants.ox.ac.uk/bol/conifers, which is the Conifer Database of
this Atlas. Data points which had no date of collection or coordinates were excluded as this infor-
mation was needed in the analysis. Each species of conifer was categorized into four main habitat
types: pine woodland/savannah, kerangas, mossy forests on ridges and summits and montane closed
canopy rainforest. For the data analysis, only the species occurring in montane closed canopy rain-
forest were selected. For these species there was the highest probability that they would no longer
be present in areas which were classified as non-forest or deforested since they were collected there.
A total of 71 samples from 10 taxa were used in the final analysis (see Table 9). Each data point
received a score for whether it occurred in remaining forest (after 2000) or a deforested area


Table 9. The predicted status of conifer species in the rainforests of Sumatera.

Species Total No. No. Estimated
No. of Predicted Predicted Decrease in
Localities Present Present Localities
(in 2000) (in 2000) (%) No. of Localities by Land Use No. of Localities in Protected Area
Forest Agri- Agriculture Total No. %
Forest Estimated Remaining
Present in PA’s
Agathis borneensis 6 1 5 83% 2 1 3 0 0 –
Dacrycarpus imbricatus var. curvulus 9 9 0 0% 9 0 0 8 8 100%
Dacrycarpus imbricatus var. imbricatus 10 4 6 60% 6 2 2 3 2 67%
Dacrydium xanthandrum 1 0 1 100% 1 0 0 0 0 –
Nageia motleyi 6 0 6 100% 1 1 4 0 0 –
Nageia wallichiana 12 1 11 92% 2 5 5 0 0 –
Podocarpus neriifolius var. neriifolius 12 4 8 67% 4 3 5 2 2 100%
examples of further research

Podocarpus teysmannii 3 0 3 100% 0 0 3 0 0 –

Sundacarpus amarus 8 3 5 63% 4 0 4 1 1 100%
Taxus wallichiana 4 3 1 25% 2 2 0 2 2 100%
Total 71 25 46 65%
481
482 examples of further research

(pre-2000). Taking into account the date at which samples were collected, a prediction was made
as to the likelihood that each data point would still represent an existing population. Further
scoring and analysis of the data points included protected areas and land use data for Sumatera.
These were then mapped using Quantum GIS (see map).
The results indicate that there is 65% likelihood that samples would no longer represent living
trees at the localities at which they have been collected. In conclusion, there is a need for further
botanical exploration in Sumatera to determine the current conservation status of these species.

Map EFR-5. Map of Sumatera showing current land use and the localities of rainforest conifers based on
herbarium collections, divided into those that have probably disappeared (red) and those that still remain
(black).

References:
Deforestation Data: http://www.Sumateranforest.org/northernSumatera.php
Land Use Data: http://www.fao.org/geonetwork/srv/en/metadata.show?id=37156
Sumatera Country Outline: http://www.gadm.org/
Aljos Farjon provided categorization of species by habitat type.
Research by Elena Shishkova, Fiona Spooner, Nicola Stanek and Lisa Wheeler from Imperial Col-
lege London studying MSc Conservation Science (February 2013).
 examples of further research 483

Modelled Conifer Diversity: a Pilot Study

Niels Raes, Denis Filer & Aljos Farjon

Whereas in previous chapters the diversity of conifers is depicted as the number of species per 1
or 0.5 degree cell, here we have developed the portrayal of patterns of diversity at a spatial resolu-
tion of 5 arc minutes, or 0.083 degrees, using species distribution modelling techniques. Species
distribution models (SDMs) relate species occurrences (georeferenced collections) with (a)biotic
predictors and interpolate the identified relationships to an area of interest; in this pilot study
the Sunda Shelf. The Sunda Shelf is located in the western part of the Malesian archipelago and is
located on a single tectonic plate (Hall, 2009b). At present, the Sunda Shelf harbours the islands
of Borneo, Sumatera, Java, Bali and the Malay Peninsula. However, during the glacial cycles of the
Pleistocene the Java Sea fell dry more than fifty times, which resulted in one continuous landmass
(Cannon et al., 2009; Woodruff, 2010). This provided ample opportunity for dispersal across the
Sunda Shelf and it is therefore regarded as one biogeographic unit.
SDMs build directly on Hutchinson’s duality, or the reciprocal correspondence between eco-
logical niche space and geographic space (Hutchinson, 1957). Plotting the ecological niche in its
reciprocal geographic space results in a geographic representation of a species’ potential distri-
bution. The presence of a species is determined by three factors that can be imagined as three
overlapping circles, each representing one factor in the ‘BAM’-framework (Soberón & Peterson,
2005). In the ‘BAM’-framework, circle ‘A’ (abiotic) represents the geographic region with appro-
priate abiotic conditions for a species to occur, and may be regarded as the geographic expression
of the fundamental abiotic niche. Circle ‘B’ (biotic) is the geographic region where the right com-
bination with interacting species occurs, which may or may not overlap extensively with ‘A’. The
intersection of ‘A’ with ‘B’ represents the geographic extent of the realized niche of the species.
The third circle ‘M’ (movement) is a representation of the geographic region that is ‘accessible’ to
the species in some ecological sense, without barriers to movement and colonization. The inter-
section of the three circles is equivalent to the observed geographic distribution of the species.
Rather than displaying diversity based solely on the verified conifer collections data, SDMs
allow us to predict the presence or absence of conifer species for areas where they have never
been collected. Such predictions and the resulting maps can be displayed with a higher spatial
resolution, depending on the resolution of the spatial predictors. Furthermore, SDMs can identify
responses to environmental gradients which may then be used to predict the impact of global
climate change, or the vulnerability of regions to alien invasion. For an extensive review on many
more advantages we refer to the literature cited above.
For this pilot study we selected all 55 conifer species occurring on the Sunda Shelf (bounded
area in Map EFR-6), which we modelled at 5 arc minute spatial resolution. We selected all collec-
tions of these species from a region extending beyond the Sunda Shelf (longitude 92° E – 127° E;
latitude 11° S – 19° N) to prevent under- and overprediction as a consequence of modelling partial
niche space (Raes, 2012). Since SDMs identify relationships between collection sites and envi-
ronmental predictors we set the lower bound for species to be modelled at five spatially unique
records. This resulted in 37 species fulfilling this requirement. For the environmental predictors
we used the Worldclim dataset (Hijmans et al., 2005), and the ISRIC soil data (Batjes, 2012) both
at 5 arc minute spatial resolution. To cater for dispersal limitation and island endemicity we
also included a categorical layer representing the islands of Borneo, Java, Sumatera, Philippines,
Sulawesi, Maluku, Lesser Sunda Islands and the Malay Peninsula/Indochina as discrete classes.
The environmental predictors were tested for correlations and those with Pearson’s r > 0.7 were
removed to prevent SDM overfitting. This resulted in 19 independent spatial predictors. Finally,
we used the non-parametric maximum entropy algorithm MaxEnt (Phillips et al., 2006; Elith
et al., 2011) to model the distributions of the 37 conifer species. The continuous MaxEnt predic-
tions were converted to discrete presence/absence predictions, and subsequently stacked result-
ing in Map EFR-6. We also aggregated the results to one degree spatial resolution (Map EFR-7) to
compare the SDM results with the count data (Map MA-3 on p. 295).
484 examples of further research

The correlation of Pearson’s r between the SDM- and count results was only 0.51, and the lin-
ear regression model explained only 0.25 percent of the variance (adjusted r 2). This is largely a
consequence of some cells having hardly any (count) collection records, but having nonetheless
predicted presence of many more species. SDMs seem to effectively fill the gaps in the count data
which are the result of opportunistic and biased plant collecting. On the other hand, the count
data include records from rare and narrow endemic species that were excluded from the models
because we have fewer than 5 records.
In conclusion, we think that the application of SDMs to produce biodiversity maps at higher
spatial resolutions has several advantages, including anticipating the impact of global climate
change on biodiversity conservation and reserve network planning. At the same time, solutions
have to be developed to model or map rare and narrow endemic species which are represented by
too few records to be modelled. Furthermore, molecular phylogenetic reconstructions combined
with SDMs pave the way to developing depictions of phylogenetic diversity patterns, which could
include information on the evolutionary and biogeographic history of a region under study.
Initial results have indicated that spatial patterns of phylogenetic diversity can be different
from taxonomic diversity patterns and therefore might have implications for future biodiversity
conservation and reserve network planning.

Map EFR-6. Conifer diversity of the Sunda Shelf (dashed line) based on 37 stacked SDMs at 5 arc minute
spatial resolution.
 examples of further research 485

Map EFR-7. Conifer diversity on the Sunda Shelf, aggregated to 1 degree spatial resolution. Points indicate
actual collections.

Species Distribution Modelling in a Phylogenetic Context Using the Conifer Database

Sarah Mathews
Using the conifer specimen data to infer species distribution models (SDM) presents exciting
opportunities to explore the interplay between ecological niche evolution and species diversi-
fication. Recent developments in the field of SDM (e.g. Elith et al., 2011) have provided tools for
using presence-only data, such as those from geo-referenced herbarium specimens, to link species
distributions with environmental characteristics of the sites they occupy. A wide variety of studies
have been enabled by these developments, including those that focus on biogeography, conserva-
tion, ecology, and invasive species. Incorporation of phylogenetic data and comparative methods
extends the utility of SDM for studies focused on understanding how ecological niches evolve and
how this has shaped species diversity (e.g. Schnitzler et al., 2012). The density and careful curation
of the Conifer Database make it particularly valuable for modelling species distributions and their
environmental correlates because the amount and accuracy of the species data mitigate potential
biases that result from limited information in herbarium and field records about prevalence of the
species in the landscape and biases in how the landscape is sampled. A few examples of potential
studies on conifers are outlined below.

Are higher rates of species turnover in Northern Hemisphere conifers associated with low rates of
climatic niche evolution?—A recently inferred age-calibrated phylogeny representing ~80% of liv-
ing conifer species revealed that node ages for conifer lineages that diversified mainly in the
Southern Hemisphere (SH) are significantly older than their counterparts in the Northern Hemi-
sphere (NH), a difference that appears to be associated with higher rates of species turnover in
NH clades (Leslie et al., 2012). By using species occurrence data in conjunction with climatic data,
486 examples of further research

e.g. from WorldClim (Hijmans et al., 2005) and phylogenetic methods, we can infer ancestral
niches and assess rates of change among climatic niches. One might predict that rates of climatic
niche evolution are low in SH clades because climatic shifts in the Southern Hemisphere have
been less dramatic than in the Northern Hemisphere. However, rates of niche evolution may be
similarly low in NH clades, and this might indicate that limited flexibility in the evolution of cli-
mate preferences has contributed to higher species turnover rates in the Northern Hemisphere.
Alternatively, if niche evolution is dynamic in NH clades, it might imply that evolution of climate
preferences has facilitated diversification in response to dramatic climatic shifts.
Within a Southern Hemisphere conifer family, do higher rates of climatic niche evolution contribute
to disparity in clade size?—Even without explicit modeling of species distributions or climatic
niches, species occurrence records facilitate tests for links between climatic range and plant
functional traits. In a recent study of diversification in Podocarpaceae, phylogenetic comparative
methods were used to test for correlated evolution of morphology and climatic range on a Podo-
carpaceae phylogenetic tree, leading to detection of interdependency between climatic range
and the presence of flattened leaves or short shoots, which enhance competitive ability under
canopies (Biffin et al., 2012). The authors also noted that genera with mostly temperate distribu-
tions were species-poor. However, most genera with tropical distributions also are small, with
none matching the success of Podocarpus (approximately 100 species). Their data suggest that
evolution of climatic tolerances may be relatively dynamic in Podocarpus and thus may have con-
tributed to its species-richness. This could be tested rigorously with the dense occurrence records
for the genus in the Conifer Database and in an explicitly SDM framework, comparing climatic
range in Podocarpus, in other tropical genera, and in temperate genera. Of course it is possible
that climatic niche is similarly conserved or dynamic across Podocarpus and the smaller genera
in Podocarpaceae, and that other ecological variables or plant functional traits have had a greater
role in differentiating clades than climate, and this can be explored by including additional types
of data in the analyses, such as soil type, elevation, architecture and breeding system.
The evolution of climatic preferences and conservation of Agathis.—Nearly two thirds (10 of 17) of
the species of Agathis (Araucariaceae) are listed as species in a threatened category on the IUCN
Red List of Threatened Species (Version 2013.2). The investigation of phylogenetic structure for
bioclimatic variables in a species phylogeny yields insights which are very relevant to conserva-
tion planning (e.g. Yesson & Culham, 2006). Specifically, by using data for 19 climate variables
from the WorldClim database (Hijmans et al., 2005) and applying SDM methods, specific cli-
mate preferences can be determined for each species. The phylogenetic distribution of climate
preferences and inference of ancestral climatic niches then identifies those lineages that would
preserve unique climatic preferences despite the extinction of individual species. Furthermore,
projections of extinction risk can be made by modeling future climatic niches based on changing
climate scenarios, in order to ask whether suitable habitats will exist in the future, and whether
they will be accessible through dispersal.

The Conifer Database Оnline

Conifers of the World Website

The Conifer Database, providing the source data for all the Atlas maps, can be consulted online
on http://herbaria.plants.ox.ac.uk/bol/conifers. With nearly 37,000 herbarium records from ca.
340 herbaria, the Conifer Database is the most comprehensive of its kind in this group of plants.
The website also includes all accepted and synonymous conifer names strengthening its value
as a research resource for anyone studying conifer biogeography and/or taxonomy. Data on the
website can be transferred to GIS systems for further analysis, including modelling. Atlas maps
can be verified and comments on these data may be returned to the authors.
 examples of further research 487

The following data are available to consult online:

■ Species: all published conifer names, authors, place and date of publication, synonymy, ety-
mology, type links and/or type notes, IUCN Red List codes, TDWG geo-coding, distribution
summary.
■ Botanical records: collector name, number, date, location text, map reference and field notes.
■ Specimens: for each botanical record specimen, herbarium location, accession number and/or
barcode, type status and determination history where available.

Data can be queried by taxonomic group (family, genus, species), geographic area (country,
major/minor region, place, map area), collector or using a custom query. The results from online
queries can be displayed in flexible tables, reports and used to verify maps.
The same database has already been used to assemble and format data for the following pub-
lications: A taxonomic monograph of Pinus in Latin America (Farjon, A. & Styles, B. T. 1997. Flora
Neotropica Monograph 75, The New York Botanical Garden); A World Checklist and Bibliography
of Conifers ( Farjon, A. 1998, 2001, Kew publishing); A Monograph of Cupressaceae and Sciadopitys
(Farjon, A. 2005. Kew Publishing); and A Handbook of the World’s Conifers (Farjon, A. 2010. Brill).

Verification of Map Points

The following example explains how to verify map points for a species on http://herbaria.plants.
ox.ac.uk/bol/conifers:

Using the home page left margin genus/species search box, type in e.g. ‘Araucaria montana’ or
sufficient letters to find the name from the list offered. You may also use the main Search menu
option to query data using different search parameters.
488 examples of further research

This leads to a query results Summary page. Here, choose Botanical Records to open a table of results.

Some of the available data processing features to help locate and verify data:

Sorting records Use one of the AZ toolbars or, for compound column sorts, drag any field on
to the Sorting order command line above the table.
Select visible columns Use the  toolbar to add or remove data columns from the list provided.
Zoom Use the  toolbar to display all non-empty data columns for the current
record.
Summary Use the  toolbar to create a list of values with record totals for the current
column.
Filter Use the  toolbar to restrict the data to match the current cell value.
Map and data verification To create a map using Google maps, select the Map option. All data points
are summarized in the right margin. Click on a coloured map cluster to zoom
further or mouse hover over an individual map marker to select and highlight
the relevant data in the display column on the right. Click on the point to open
its data text box. You can also click on the Zoom button for any record in the
map data summary to locate and zoom to that point. Note that map points are
to the nearest minute and that the google maps terrain will not be precise.

Map EFR-8. The distribution of the species Araucaria montana in New Caledonia plotted using Google
Maps. Data points are summarized in right-side pane. Mouse hover over individual points to highlight map
point details in the data margin. Click on cluster points to zoom. Alternatively, click on the Zoom button
in the data margin to locate and zoom a point on the map.
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 Tables and Indices

Table 10. Countries with numbers of taxa, endemics and IUCN Red List categories of threat.

The list of countries includes several overseas dependencies or territories because of their distinct biogeo­
graphy; in no case is any sovereign status expressed or implied and neither do we claim that the list is
complete. Taxa are species and infraspecific ranks. IUCN Red List categories used are: Critically Endangered
(CR), Endangered (EN), Vulnerable (VU), Near Threatened (NT), Least Concern (LC), Data Deficient (DD)
and Not Evaluated (NE). The Red List status is global for all taxa, not regional (e.g. a country in cases where
a taxon occurs in more than one country); the figures therefore indicate how many globally threatened
conifer taxa each country has within its borders. The categories of threat are CR, EN and VU.

Country Total taxa Endemic CR EN VU NT and LC Notes

Afghanistan 11 0 0 1 0 10
Albania 14 0 0 0 0 13 1 NE
Algeria 15 2 1 2 0 12
American Samoa 0
Andorra 4 0 0 0 0 4
Angola 1 0 0 0 0 1
Antarctica 0
Antigua and Barbuda 0
Argentina 9 0 1 2 1 5
Armenia 7 0 0 0 0 7
Aruba 0
Australia 40 39 2 2 6 31
Austria 16 0 0 1 0 15
Azerbaijan 9 0 0 0 0 8 1 DD
Bahamas 2 0 0 0 2 0
Bahrain 0
Bangladesh 2 0 0 0 0 2
Barbados 1 0 1 0 0 0
Belarus 4 0 0 0 0 4
Belgium 2 0 0 0 0 2
Belize 4 0 0 0 1 3
Benin 0
Bermuda 1 1 1 0 0 0
Bhutan 15 0 0 2 1 13
Bolivia 10 1 0 0 1 8 1 NE
Bosnia and Herzegovina 11 0 0 1 0 10
Botswana 0
Brazil 10 2 2 2 0 5 1 DD
British Virgin Islands 0
Brunei 12 0 0 2 1 9
Bulgaria 14 0 0 0 0 13 1 NE
Burkina Faso 0
Burundi 2 0 0 0 0 2
Cambodia 8 0 0 0 1 7
Cameroon 2 0 0 0 1 1
Canada 42 2 0 1 0 41
Cape Verde Islands 0
Cayman Islands 0
Central African Republic 0
 tables and indices 495

Table (cont.)
Country Total taxa Endemic CR EN VU NT and LC Notes
Chad 0
Chile 9 1 0 2 3 4
China 190 112 7 23 35 115 10 DD
Christmas Island 0
Cocos (Keeling) Islands 0
Colombia 8 0 0 0 2 5 1 NE
Comoros 0
Congo 0
Congo Democratic Republic 2 0 0 0 0 4
Cook Islands 0
Costa Rica 4 2 1 1 0 1 1 NE
Cote d’Ivoire 0
Croatia 15 1 0 1 0 14
Cuba 7 6 2 1 2 2
Cyprus 8 1 0 0 1 7
Czech Republic 10 0 0 1 0 9
Denmark 2 0 0 0 0 2
Djibouti 0
Dominica 1 0 0 0 0 1
Dominican Republic 6 3 1 5 0 0
East Timor 2 0 0 0 0 2
Ecuador 9 0 0 0 1 5 1 NE
Egypt 1 0 0 0 0 1
El Salvador 12 0 0 3 2 6 1 NE
Equatorial Guinea 0
Eritrea 1 0 0 0 0 1
Estonia 4 0 0 0 0 4
Ethiopia 2 0 0 0 0 2
Falkland Islands 0 plantations
Faroe Islands 1 0 0 0 0 1
Fiji 9 3 1 2 0 6
Finland 5 0 0 0 0 5
France 26 0 1 1 0 24
French Guiana 0
French Polynesia 0
Gabon 0
Gambia 0
Georgia 15 0 0 0 1 13 1 DD
Germany 12 0 0 1 0 11
Ghana 0
Gibraltar 1 0 0 0 0 1
Greece 22 1 0 0 0 21 1 NE
Greenland 1 0 0 0 0 1
Grenada 0
Guadeloupe 1 0 0 0 0 1
Guam 0
Guatemala 22 0 0 6 2 13 1 NE
Guinea 0
Guinea-Bissau 0
Guyana 3 0 0 0 0 3
Haiti 5 0 1 3 1 0
Honduras 14 0 0 2 2 9 1 NE
Hungary 2 0 0 0 0 2
Iceland 1 0 0 0 0 1
India 31 2 0 3 2 26
Indonesia 56 4 0 5 7 43 1 DD
Iran 11 0 0 0 0 10 1 DD
Iraq 3 0 0 0 0 2 1 DD
496 tables and indices

Table (cont.)
Country Total taxa Endemic CR EN VU NT and LC Notes
Ireland 3 0 0 0 0 3
Israel 3 0 0 0 0 3
Italy 21 0 1 0 0 20
Jamaica 3 2 1 1 1 0
Japan 54 36 1 5 2 46 1 DD
Jordan 2 0 0 0 0 2
Kazakhstan 10 0 0 0 1 9
Kenya 4 0 0 1 0 3
Kirgyzstan 8 0 0 0 1 7
Kuwait 0
Lao PDR 16 0 1 1 4 10
Latvia 2 0 0 0 0 2
Lebanon 8 0 0 0 1 7
Lesotho 0
Liberia 0
Libya 4 0 0 0 0 4
Liechtenstein 11 0 0 0 0 11
Lithuania 2 0 0 0 0 2
Luxembourg 2 0 0 0 0 2
Macaronesia 5 3 0 1 1 3
Macau 0
Macedonia 13 0 0 0 0 13
Madagascar 7 7 1 4 0 1 1 DD
Malawi 7 1 1 1 0 5
Malaysia 42 16 0 10 9 22 1 DD
Maldives 0
Mali 0
Malta 2 0 0 0 0 2
Marshall Islands 0
Martinique 0
Mauritania 0
Mauritius 0 plantations
Mexico 110 45 2 21 14 70 1 DD, 2 NE
Micronesia 0
Moldova 0 plantations
Monaco 3 0 0 0 0 3
Mongolia 12 0 0 0 0 12
Montenegro 10 0 0 0 0 10
Montserrat 0
Morocco 14 3 0 3 0 11
Mozambique 3 0 0 0 0 3
Myanmar [Burma] 21 0 0 2 3 16
Namibia 0
Nauru 0
Nepal 21 0 0 2 2 17
Netherlands 3 0 0 0 0 2
Netherlands Antilles 0
New Caledonia 43 43 4 13 7 18 1 NE
New Zealand 20 20 0 0 1 19
Nicaragua 7 0 0 0 1 5 1 NE
Niger 0
Nigeria 1 0 0 0 0 1
Niue 0
Norfolk Island 1 1 0 0 1 0
North Korea 17 0 0 1 2 14
Northern Mariana Islands 0
Norway 7 0 0 0 0 7
 tables and indices 497

Table (cont.)
Country Total taxa Endemic CR EN VU NT and LC Notes
Oman 1 0 0 0 0 1
Pakistan 13 0 0 1 0 12
Palau 0
Palestine 2 0 0 0 0 2
Panama 3 0 0 0 0 2 1 NE
Papua New Guinea 36 2 0 0 1 35
Paraguay 0 plantations
Peru 9 0 0 1 2 4 1 NE
Philippines 25 4 1 4 4 15 1 DD
Pitcairn Islands 0
Poland 11 1 0 1 1 9
Portugal 11 0 0 0 0 11
Puerto Rico (USA) 1 0 0 0 0 1
Qatar 0
Réunion 0
Romania 10 0 0 0 0 10
Russia 39 0 0 0 2 36 1 NE
Rwanda 2 0 0 0 0 2
Saint Kitts and Nevis 1 0 0 0 0 1
Saint Lucia 2 1 1 0 0 1
Saint Vincent/Grenadines 0
Samoa 0
San Marino 0
São Tomé and Príncipe 1 1 0 0 1 0
Saudi Arabia 2 0 0 0 0 2
Senegal 0
Serbia 6 0 0 1 0 5
Seychelles 0
Sierra Leone 0
Singapore 1 0 0 0 1 0
Slovakia 7 0 0 1 0 6
Slovenia 5 0 0 0 0 5
Solomon Islands 11 1 0 2 0 9
Somalia 1 0 0 0 0 1
South Africa 8 3 1 1 1 5
South Korea 18 0 0 1 1 15 1 NE
South Sudan 0
Spain 21 1 0 1 0 20
Sri Lanka 0 plantations
Sudan 2 0 0 0 0 2
Surinam 0
Svalbard and Jan Mayen Island 0
Swaziland 1 0 0 0 0 1
Sweden 6 0 0 0 0 6
Switzerland 13 0 0 1 0 12
Syria 9 0 0 0 1 8
Taiwan 29 12 0 8 6 14 1 DD
Tajikistan 3 0 0 0 0 3
Tanzania 6 0 0 2 0 4
Thailand 10 0 0 1 2 7
Togo 0
Tonga 1 1 0 0 1 0
Trinidad and Tobago 2 1 0 0 0 2
Tunisia 7 0 0 1 0 6
Turkey 26 5 0 0 1 23 2 NE
Turkmenistan 4 0 0 0 0 4
Turks and Caicos Islands 1 0 0 0 1 0
Tuvalu 0
498 tables and indices

Table (cont.)
Country Total taxa Endemic CR EN VU NT and LC Notes
United States of America 125 82 7 10 6 101 1 NE
Uganda 4 0 0 0 0 4
Ukraine 16 0 0 0 1 15
United Arab Emirates 0
United Kingdom 4 0 0 0 0 4
Uruguay 0 plantations
Uzbekistan 5 0 0 0 1 4
Vanuatu 8 1 0 1 0 7
Venezuela 15 1 0 1 2 11 1 NE
Viet Nam 33 7 1 6 10 16
Virgin Islands (US) 0
Western Sahara 0
Windward Islands 1 0 0 0 0 1
Yemen 1 0 0 0 0 1
Zambia 2 0 0 0 0 2
Zimbabwe 5 0 0 1 0 4

Family Index

Maps of the families Cephalotaxaceae, Phyllocladaceae and Sciadopityaceae are equivalent to those of their
eponymous single genera (see genus index) under which a larger map is given.

Family Map number Page

Araucariaceae GTC-11, GTC-17 16, 22
Cephalotaxaceae GTC-8, MAJ-108 15, 242
Cupressaceae GTC-5, GTC-13 14, 17
Phyllocladaceae GTC-12 16
Pinaceae GTC-6, GTC-14 14, 19
Podocarpaceae GTC-10, GTC-16 15, 21
Sciadopityaceae GTC-9 15
Taxaceae GTC-7, GTC-15 15, 20

Genus Index

Maps of monospecific genera are equivalent to their single species and are presented as a species map (see
species index); maps of genera with 2–3 species are also omitted if their distribution can be easily obtained
from the species maps.

Genus Family Map number Page

Abies Pinaceae GTC-21 28
Acmopyle Podocarpaceae ‒
Actinostrobus Cupressaceae AU-12 352
Afrocarpus Podocarpaceae AF-11 465
Agathis Araucariaceae MA-66 332
Amentotaxus Taxaceae MAJ-115 246
Araucaria Araucariaceae AU-74 391
Athrotaxis Cupressaceae ‒
Austrocedrus Cupressaceae ‒
Austrotaxus Taxaceae ‒
Callitris Cupressaceae AU-16 354
Calocedrus Cupressaceae MAJ-105 241
 tables and indices 499

Table (cont.)
Genus Family Map number Page
Cathaya Pinaceae ‒
Cedrus Pinaceae EM-26 157
Cephalotaxus Cephalotaxaceae MAJ-108 242
Chamaecyparis Cupressaceae MAJ-181 287
Cryptomeria Cupressaceae ‒
Cunninghamia Cupressaceae ‒
Cupressus Cupressaceae NA-70 76
Dacrycarpus Podocarpaceae MA-43 316
Dacrydium Podocarpaceae MA-31 309
Diselma Cupressaceae ‒
Falcatifolium Podocarpaceae MA-50 320
Fitzroya Cupressaceae ‒
Fokienia Cupressaceae ‒
Glyptostrobus Cupressaceae ‒
Halocarpus Podocarpaceae ‒
Juniperus Cupressaceae GTC-22 29
Keteleeria Pinaceae MAJ-68 218
Lagarostrobos Podocarpaceae ‒
Larix Pinaceae GTC-20 26
Lepidothamnus Podocarpaceae ‒
Libocedrus Cupressaceae AU-48 375
Manoao Podocarpaceae ‒
Metasequoia Cupressaceae ‒
Microbiota Cupressaceae ‒
Microcachrys Podocarpaceae ‒
Nageia Podocarpaceae MA-54 323
Neocallitropsis Cupressaceae ‒
Nothotsuga Pinaceae ‒
Papuacedrus Cupressaceae ‒
Parasitaxus Podocarpaceae ‒
Pherosphaera Podocarpaceae ‒
Phyllocladus Phyllocladaceae AU-64 384
Picea Pinaceae GTC-19 25
Pilgerodendron Cupressaceae AU-48 375
Pinus Pinaceae GTC-18 23
Platycladus Cupressaceae ‒
Podocarpus Podocarpaceae GTC-24 32
Prumnopitys Podocarpaceae SA-5 432
Pseudolarix Pinaceae ‒
Pseudotaxus Taxaceae ‒
Pseudotsuga Pinaceae NA-44 59
Retrophyllum Podocarpaceae AU-100 410
Saxegothaea Podocarpaceae ‒
Sciadopitys Sciadopityaceae ‒
Sequoia Cupressaceae ‒
Sequoiadendron Cupressaceae ‒
Sundacarpus Podocarpaceae ‒
Taiwania Cupressaceae ‒
Taxodium Cupressaceae ‒
Taxus Taxaceae GTC-23 31
Tetraclinis Cupressaceae ‒
Thuja Cupressaceae MAJ-179 286
Thujopsis Cupressaceae ‒
Torreya Taxaceae MAJ-118 247
Tsuga Pinaceae MAJ-167 279
Widdringtonia Cupressaceae AF-6 463
Wollemia Araucariaceae ‒
Xanthocyparis Cupressaceae ‒
500 tables and indices

Species Index

Only species (binomials) are listed; if subspecies or varieties are recognized, they are in nearly all cases
presented in the map of the species. The number of degree squares is based on the number of map points
(herbarium collections from different localities) that fall in different degree squares. This figure gives an
indication of the extent of occurrence (EOO) as well as the area of occupancy (AOO) of a species, but is
dependent on the representation of herbarium specimens in the Conifer Database relative to the actual
range and abundance of the species. Very widespread and ubiquitous species in the northern boreal forests
of Canada and Siberia are under-represented and are marked with *. The IUCN Red List status is for the
species; if subspecies or varieties are recognized these may have a different status. This information can be
accessed at http://www.iucnredlist.org/

Species Map number Page Photo page Degree squares IUCN Red List
Abies alba EM-20 153 154 69 LC
Abies amabilis NA-60 68 29 33 LC
Abies balsamea NA-55 65 92* LC
Abies beshanzuensis MAJ-82 225 226 1 CR
Abies bracteata NA-63 69 70 3 NT
Abies cephalonica EM-22 155 8 LC
Abies chensiensis MAJ-73 221 21 LC
Abies cilicica MAJ-6 177 14 NT
Abies concolor NA-61 68 60 LC
Abies delavayi MAJ-75 222 18 LC
Abies densa MAJ-24 189 9 LC
Abies durangensis MCW-35 111 15 LC
Abies fabri MAJ-74 221 8 VU
Abies fanjingshanensis MAJ-82 225 1 EN
Abies fargesii MAJ-76 222 40 LC
Abies firma MAJ-170 280 28 LC
Abies forrestii MAJ-77 223 223 41 LC
Abies fraseri NA-62 69 4 EN
Abies grandis NA-57 66 48 LC
Abies guatemalensis MCW-39 113 15 EN
Abies hickelii MCW-37 112 5 EN
Abies hidalgensis MCW-35 111 1 VU
Abies holophylla MAJ-80 224 17 NT
Abies homolepis MAJ-173 281 11 NT
Abies kawakamii MAJ-129 253 254 3 NT
Abies koreana MAJ-147 266 3 EN
Abies lasiocarpa NA-56 65 65 107 LC
Abies magnifica NA-59 67 67 28 LC
Abies mariesii MAJ-171 281 13 LC
Abies nebrodensis EM-23 155 1 CR
Abies nephrolepis MAJ-81 225 21 LC
Abies nordmanniana MAJ-7 177 178 23 LC
Abies numidica EM-25 156 1 CR
Abies pindrow MAJ-25 190 190 16 LC
Abies pinsapo EM-24 155 3 EN
Abies procera NA-58 66 10 LC
Abies recurvata MAJ-79 224 28 VU
Abies religiosa MCW-38 113 113 20 LC
Abies sachalinensis MAJ-174 282 282 14 LC
Abies sibirica MAJ-13 181 48* LC
Abies spectabilis MAJ-25 190 14 NT
Abies squamata MAJ-78 223 17 VU
Abies veitchii MAJ-172 281 13 LC
Abies vejarii MCW-36 112 9 NT
Abies ×borisii-regis EM-21 154 9 NE
Abies yuanbaoshanensis MAJ-82 225 1 CR
 tables and indices 501

Table (cont.)
Species Map number Page Photo page Degree squares IUCN Red List
Abies ziyuanensis MAJ-82 225 3 EN
Acmopyle pancheri AU-98 408 408 5 NT
Acmopyle sahniana AU-119 423 2 CR
Actinostrobus acuminatus AU-15 353 4 NT
Actinostrobus arenarius AU-14 353 353 10 LC
Actinostrobus pyramidalis AU-13 352 10 LC
Afrocarpus dawei AF-15 467 7 NT
Afrocarpus falcatus AF-14 467 24 LC
Afrocarpus gracilior AF-12 466 466 33 LC
Afrocarpus mannii AF-16 468 1 VU
Afrocarpus usambarensis AF-13 467 9 EN
Agathis atropurpurea AU-7 348 2 NT
Agathis australis AU-47 374 373, 374 9 NT
Agathis borneensis MA-68 333 36 EN
Agathis dammara MA-69 334 36 VU
Agathis flavescens MA-70 335 1 VU
Agathis kinabaluensis MA-70 335 2 EN
Agathis labillardierei MA-72 337 22 NT
Agathis lanceolata AU-71 389 4 VU
Agathis lenticula MA-71 336 336 5 VU
Agathis macrophylla AU-116 421 10 EN
Agathis microstachya AU-6 347 347 1 NT
Agathis montana AU-73 390 1 NT
Agathis moorei AU-70 389 6 VU
Agathis orbicula MA-67 333 4 EN
Agathis ovata AU-72 390 390 3 EN
Agathis robusta AU-8 348 13 LC
Agathis silbae AU-117 422 2 NT
Amentotaxus argotaenia MAJ-116 246 246 37 NT
Amentotaxus assamica MAJ-34 196 2 EN
Amentotaxus formosana MAJ-133 255 1 EN
Amentotaxus hatuyenensis MAJ-143 263 1 EN
Amentotaxus poilanei MAJ-143 263 1 VU
Amentotaxus yunnanensis MAJ-117 247 7 VU
Araucaria angustifolia SA-23 443 444 20 CR
Araucaria araucana SA-30 449 450 7 EN
Araucaria bernieri AU-80 395 396 2 VU
Araucaria bidwillii AU-9 349 350 4 LC
Araucaria biramulata AU-81 396 4 VU
Araucaria columnaris AU-76 393 393 6 LC
Araucaria cunninghamii AU-10 350 350 51 LC
Araucaria heterophylla AU-118 422 422 1 VU
Araucaria humboldtensis AU-87 400 2 EN
Araucaria hunsteinii MA-65 331 5 NT
Araucaria laubenfelsii AU-82 397 5 NT
Araucaria luxurians AU-83 397 397 5 EN
Araucaria montana AU-78 394 6 VU
Araucaria muelleri AU-86 399 3 EN
Araucaria nemorosa AU-88 400 400 2 CR
Araucaria rulei AU-85 399 399 5 EN
Araucaria schmidii AU-77 394 1 VU
Araucaria scopulorum AU-84 398 3 EN
Araucaria subulata AU-79 395 2 NT
Athrotaxis cupressoides AU-38 367 5 VU
Athrotaxis selaginoides AU-39 367 367 6 VU
Athrotaxis ×laxifolia AU-40 368 4 VU
Austrocedrus chilensis SA-31 450 451 23 NT
Austrotaxus spicata AU-94 405 3 NT
502 tables and indices

Table (cont.)
Species Map number Page Photo page Degree squares IUCN Red List
Callitris baileyi AU-24 358 5 NT
Callitris canescens AU-20 356 356 23 LC
Callitris columellaris AU-17 354 346, 355 80 LC
Callitris drummondii AU-22 357 7 NT
Callitris endlicheri AU-23 357 28 LC
Callitris macleayana AU-26 359 359 9 LC
Callitris monticola AU-25 359 4 EN
Callitris muelleri AU-24 358 358 4 NT
Callitris neocaledonica AU-92 403 2 NT
Callitris oblonga AU-28 361 7 VU
Callitris preissii AU-18 355 48 LC
Callitris rhomboidea AU-27 360 31 LC
Callitris roei AU-21 357 357 9 EN
Callitris sulcata AU-91 402 2 EN
Callitris verrucosa AU-19 356 33 LC
Calocedrus decurrens NA-90 86 31 LC
Calocedrus formosana MAJ-131 254 2 EN
Calocedrus macrolepis MAJ-106 241 32 NT
Calocedrus rupestris MAJ-141 261 10 EN
Cathaya argyrophylla MAJ-47 207 8 VU
Cedrus atlantica EM-27 157 11 EN
Cedrus deodara MAJ-15 183 184 19 LC
Cedrus libani EM-28 157 158 17 VU
Cephalotaxus fortunei MAJ-109 242 243 90 LC
Cephalotaxus hainanensis MAJ-113 244 3 EN
Cephalotaxus harringtonii MAJ-188 290 54 LC
Cephalotaxus lanceolata MAJ-114 245 1 EN
Cephalotaxus latifolia MAJ-112 244 10 NT
Cephalotaxus mannii MAJ-142 262 32 VU
Cephalotaxus oliveri MAJ-110 243 16 VU
Cephalotaxus sinensis MAJ-111 243 60 LC
Chamaecyparis formosensis MAJ-132 255 255 4 EN
Chamaecyparis lawsoniana NA-68 74 10 NT
Chamaecyparis obtusa MAJ-183–184 288 255 33 NT
Chamaecyparis pisifera MAJ-182 287 17 LC
Chamaecyparis thyoides NA-67 73 30 LC
Cryptomeria japonica MAJ-177 284 285 17 NT
Cunninghamia konishii MAJ-88 230 7 EN
Cunninghamia lanceolata MAJ-87 229 229 74 LC
Cupressus arizonica NA-77 79 33 LC
Cupressus bakeri NA-73 77 7 EN
Cupressus cashmeriana MAJ-26 191 7 NT
Cupressus chengiana MAJ-94 234 8 VU
Cupressus duclouxiana MAJ-95 235 235 13 DD
Cupressus dupreziana EM-30 159 160 3 EN
Cupressus funebris MAJ-96 236 31 DD
Cupressus goveniana NA-74 77 5 EN
Cupressus guadalupensis NA-76 78 5 EN
Cupressus lusitanica MCW-42 115 31 LC
Cupressus macnabiana NA-72 77 8 LC
Cupressus macrocarpa NA-75 78 78 1 VU
Cupressus sargentii NA-71 76 10 VU
Cupressus sempervirens EM-29 159 29 LC
Cupressus torulosa MAJ-27 191 22 LC
Dacrycarpus cinctus MA-48 319 19 LC
Dacrycarpus compactus MA-46 318 18 LC
Dacrycarpus cumingii MA-45 318 16 LC
Dacrycarpus dacrydioides AU-58 381 373 29 LC
 tables and indices 503

Table (cont.)
Species Map number Page Photo page Degree squares IUCN Red List
Dacrycarpus expansus MA-47 318 7 LC
Dacrycarpus imbricatus MA-44 317 317 116 LC
Dacrycarpus kinabaluensis MA-63 329 329 1 LC
Dacrycarpus steupii MA-49 319 18 NT
Dacrycarpus vieillardii AU-99 409 4 LC
Dacrydium araucarioides AU-112 416 416 3 LC
Dacrydium balansae AU-110 415 5 LC
Dacrydium beccarii MA-33 311 48 LC
Dacrydium comosum MA-36 312 313 3 EN
Dacrydium cornwallianum MA-39 314 7 LC
Dacrydium cupressinum AU-57 380 380 35 LC
Dacrydium elatum MA-35 312 39 LC
Dacrydium ericoides MA-37 313 4 LC
Dacrydium gibbsiae MA-60 328 1 LC
Dacrydium gracile MA-36 312 3 NT
Dacrydium guillauminii AU-113 417 417 1 CR
Dacrydium leptophyllum MA-42 315 1 VU
Dacrydium lycopodioides AU-111 416 2 NT
Dacrydium magnum MA-42 315 4 NT
Dacrydium medium MA-37 313 6 VU
Dacrydium nausoriense AU-120 424 3 EN
Dacrydium nidulum MA-41 315 31 LC
Dacrydium novoguineense MA-40 314 13 LC
Dacrydium pectinatum MA-34 311 27 EN
Dacrydium spathoides MA-38 313 1 NT
Dacrydium ×suprinii AU-114 418 1 NE
Dacrydium xanthandrum MA-32 310 310 23 LC
Diselma archeri AU-45 370 371 5 LC
Falcatifolium angustum MA-51 321 2 EN
Falcatifolium falciforme MA-53 322 322 18 NT
Falcatifolium gruezoi MA-53 322 10 NT
Falcatifolium papuanum MA-52 321 5 LC
Falcatifolium sleumeri MA-52 321 1 NT
Falcatifolium taxoides AU-96 407 6 LC
Fitzroya cupressoides SA-32 451 10 EN
Fokienia hodginsii MAJ-93 234 216 47 VU
Glyptostrobus pensilis MAJ-91 232 17 CR
Halocarpus bidwillii AU-60 382 382 24 LC
Halocarpus biformis AU-59 382 31 LC
Halocarpus kirkii AU-59 382 7 VU
Juniperus angosturana MCW-45 117 8 VU
Juniperus arizonica NA-87 84 13 LC
Juniperus ashei NA-85 84 29 LC
Juniperus barbadensis MCW-65 130 130 15 VU
Juniperus bermudiana MCW-64 129 1 CR
Juniperus blancoi MCW-51 119 9 NT
Juniperus brevifolia EM-39 167 4 VU
Juniperus californica NA-84 83 83 22 LC
Juniperus cedrus EM-40 167 168 4 EN
Juniperus chinensis MAJ-98 237 74 LC
Juniperus coahuilensis MCW-46 117 12 LC
Juniperus comitana MCW-53 121 6 EN
Juniperus communis EM-31–32 160, 161 162, 178 492* LC
Juniperus convallium MAJ-99 238 29 LC
Juniperus deppeana MCW-52 120  30, 121 44 LC
Juniperus drupacea EM-38 166 14 LC
Juniperus durangensis MCW-49 118 9 LC
504 tables and indices

Table (cont.)
Species Map number Page Photo page Degree squares IUCN Red List
Juniperus excelsa MAJ-8 178 109 LC
Juniperus flaccida MCW-44 117 117 32 LC
Juniperus foetidissima EM-37 165 166 46 LC
Juniperus formosana MAJ-104 240 71 LC
Juniperus gamboana MCW-54 121 3 EN
Juniperus gracilior MCW-66 131 6 EN
Juniperus horizontalis NA-80 81 113* LC
Juniperus indica MAJ-29 192 43 LC
Juniperus jaliscana MCW-48 118 2 EN
Juniperus komarovii MAJ-102 239 9 NT
Juniperus monosperma NA-83 83 30 32 LC
Juniperus monticola MCW-50 119 11 LC
Juniperus occidentalis NA-88 85 85 52 LC
Juniperus osteosperma NA-82 82 82 56 LC
Juniperus oxycedrus EM-34 163 162 LC
Juniperus phoenicea EM-35 164 116 LC
Juniperus pinchotii NA-86 84 32 LC
Juniperus pingii MAJ-103 239 240 49 LC
Juniperus procera AF-5 462 462 45 LC
Juniperus procumbens MAJ-187 290 4 LC
Juniperus przewalskii MAJ-102 239 20 LC
Juniperus pseudosabina MAJ-14 182 182 55 LC
Juniperus recurva MAJ-30 193 39 LC
Juniperus rigida MAJ-185 288 62 LC
Juniperus sabina EM-33 162 105* LC
Juniperus saltillensis MCW-47 118 9 EN
Juniperus saltuaria MAJ-101 238 24 LC
Juniperus saxicola MCW-67 131 2 CR
Juniperus scopulorum NA-81 81 72 LC
Juniperus semiglobosa MAJ-31 193 194 49 LC
Juniperus squamata MAJ-28 192 58 LC
Juniperus standleyi MCW-55 122 2 EN
Juniperus taxifolia MAJ-186 289 289 7 NT
Juniperus thurifera EM-36 165 26 LC
Juniperus tibetica MAJ-100 238 47 VU
Juniperus virginiana NA-89 86 109 LC
Keteleeria davidiana MAJ-69 219 42 LC
Keteleeria evelyniana MAJ-71 220 26 VU
Keteleeria fortunei MAJ-70 219 19 NT
Lagarostrobos franklinii AU-42 369 5 LC
Larix decidua EM-19 152 33 LC
Larix gmelinii MAJ-146 266 71* LC
Larix griffithii MAJ-21 188 29 LC
Larix kaempferi MAJ-166 278 7 LC
Larix laricina NA-47 61 131* LC
Larix lyallii NA-48 61 21 LC
Larix mastersiana MAJ-62 214 4 EN
Larix occidentalis NA-49 62 37 LC
Larix potaninii MAJ-22, 63 188, 215 33 LC
Larix sibirica MAJ-12 181 27 60* LC
Larix ×czekanowskii ‒ ‒ NE
Lepidothamnus fonkii SA-34 453 19 LC
Lepidothamnus intermedius AU-62 383 23 LC
Lepidothamnus laxifolius AU-63 383 23 LC
Libocedrus austrocaledonica AU-89 401 3 NT
Libocedrus bidwillii AU-49 375 376 18 NT
Libocedrus chevalieri AU-90 402 2 CR
 tables and indices 505

Table (cont.)
Species Map number Page Photo page Degree squares IUCN Red List
Libocedrus plumosa AU-50 376 13 NT
Libocedrus yateensis AU-90 402 402 2 EN
Manoao colensoi AU-61 383 22 LC
Metasequoia glyptostroboides MAJ-90 231 1 EN
Microbiota decussata MAJ-148 267 5 LC
Microcachrys tetragona AU-44 370 370 5 LC
Nageia fleuryi MAJ-139 260 21 NT
Nageia maxima MA-57 324 2 EN
Nageia motleyi MA-56 324 18 VU
Nageia nagi MAJ-83 226 227 43 NT
Nageia wallichiana MA-55 323 109 LC
Neocallitropsis pancheri AU-93 404 403 2 EN
Nothotsuga longibracteata MAJ-67 217 218 17 NT
Papuacedrus papuana MA-73 337 31 LC
Parasitaxus usta AU-95 406 406 4 VU
Pherosphaera fitzgeraldii AU-36 365 1 CR
Pherosphaera hookeriana AU-43 369 3 NT
Phyllocladus aspleniifolius AU-41 368 11 LC
Phyllocladus hypophyllus MA-64 330 64 LC
Phyllocladus toatoa AU-65 385 11 LC
Phyllocladus trichomanoides AU-66–67 385, 386 386 53 LC
Picea abies EM-17 151 152 149 LC
Picea alcoquiana MAJ-164 276 3 NT
Picea asperata MAJ-49 208 26 VU
Picea aurantiaca MAJ-50 209 2 EN
Picea brachytyla MAJ-52 209 210 1 VU
Picea breweriana NA-43 58 5 VU
Picea chihuahuana MCW-41 114 4 EN
Picea crassifolia MAJ-56 211 28 LC
Picea engelmannii NA-39 57 80 LC
Picea farreri MAJ-138 259 2 VU
Picea glauca NA-38 56 26, 56 123* LC
Picea glehnii MAJ-160 274 10 LC
Picea jezoensis MAJ-159 274 40 LC
Picea koraiensis MAJ-60 213 14 LC
Picea koyamae MAJ-162 275 2 CR
Picea likiangensis MAJ-54 210 42 VU
Picea linzhiensis MAJ-55 211 9 LC
Picea mariana NA-37 55 140* LC
Picea martinezii MCW-40 114 3 EN
Picea maximowiczii MAJ-161 275 4 EN
Picea meyeri MAJ-57 211 13 NT
Picea morrisonicola MAJ-128 253 3 VU
Picea neoveitchii MAJ-59 212 7 CR
Picea obovata MAJ-10 180 41* LC
Picea omorika EM-18 152 2 EN
Picea orientalis MAJ-5 176 13 LC
Picea pungens NA-42 58 38 LC
Picea purpurea MAJ-53 210 16 NT
Picea retroflexa MAJ-51 209 7 EN
Picea rubens NA-40 57 47 LC
Picea schrenkiana MAJ-11 180 180 33 LC
Picea sitchensis NA-41 57 54 LC
Picea smithiana MAJ-20 187 13 LC
Picea spinulosa MAJ-20 187 9 LC
Picea torano MAJ-163 276 10 VU
Picea wilsonii MAJ-58 212 31 LC
Picea ×fennica ‒ ‒ NE
506 tables and indices

Table (cont.)
Species Map number Page Photo page Degree squares IUCN Red List
Picea ×lutzii ‒ ‒ NE
Pilgerodendron uviferum SA-33 452 453 30 VU
Pinus albicaulis NA-17 45 67 EN
Pinus amamiana MAJ-157 273 1 EN
Pinus aristata NA-15 43 43 13 LC
Pinus arizonica MCW-21 103 102 LC
Pinus armandii MAJ-38 202 62 LC
Pinus attenuata NA-22 49 28 LC
Pinus ayacahuite MCW-26 106 31 LC
Pinus balfouriana NA-15 43 43 7 NT
Pinus banksiana NA-6 39 89* LC
Pinus bhutanica MAJ-19 186 187 7 LC
Pinus brutia EM-15 149 53 LC
Pinus bungeana MAJ-36 201 24 LC
Pinus canariensis EM-16 150 150 5 LC
Pinus caribaea MCW-71 133 134 39 LC
Pinus cembra EM-13 148 22 LC
Pinus cembroides MCW-27 106 107 79 LC
Pinus clausa NA-32 53 16 LC
Pinus contorta NA-20 47 116 LC
Pinus coulteri NA-25 50 17 NT
Pinus cubensis MCW-70 133 6 LC
Pinus culminicola MCW-28 107 2 EN
Pinus dalatensis MAJ-137 259 8 VU
Pinus densata MAJ-44 205 30 LC
Pinus densiflora MAJ-155 271 272 59 LC
Pinus devoniana MCW-19 102 102 32 LC
Pinus douglasiana MCW-13 99 19 LC
Pinus durangensis MCW-25 105 20 NT
Pinus echinata NA-27 51 62 LC
Pinus edulis NA-12 42 47 LC
Pinus elliottii NA-31 53 25 LC
Pinus engelmannii MCW-22 104 28 LC
Pinus fenzeliana MAJ-39 203 203 24 NT
Pinus flexilis NA-16 44 44 76 LC
Pinus gerardiana MAJ-16 184 185 14 NT
Pinus glabra NA-33 53 25 LC
Pinus greggii MCW-17 101 9 VU
Pinus halepensis EM-14 149 72 LC
Pinus hartwegii MCW-10 97 98 28 LC
Pinus heldreichii EM-11 147 20 LC
Pinus henryi MAJ-45 206 12 NT
Pinus herrerae MCW-24 105 19 LC
Pinus hwangshanensis MAJ-46 206 206 30 LC
Pinus jaliscana MCW-14 99 3 NT
Pinus jeffreyi NA-11 42 31 LC
Pinus kesiya MAJ-135 257 41 LC
Pinus koraiensis MAJ-145 265 28 LC
Pinus krempfii MAJ-136 258 258 2 VU
Pinus lambertiana NA-10 41 34 LC
Pinus latteri MAJ-134 256 257 28 NT
Pinus lawsonii MCW-18 101 18 LC
Pinus leiophylla MCW-16 100 64 LC
Pinus longaeva NA-15 43 20 LC
Pinus luchuensis MAJ-156 272 7 LC
Pinus lumholtzii MCW-18 101 21 NT
Pinus luzmariae MCW-8 96 11 LC
Pinus massoniana MAJ-41 204 82 LC
 tables and indices 507

Table (cont.)
Species Map number Page Photo page Degree squares IUCN Red List
Pinus maximartinezii MCW-33 110 110 3 EN
Pinus maximinoi MCW-11 98 43 LC
Pinus merkusii MA-74 338 9 VU
Pinus monophylla NA-14 42 42 43 LC
Pinus montezumae MCW-20 103 103 42 LC
Pinus monticola NA-9 41 44 NT
Pinus morrisonicola MAJ-126 252 4 NT
Pinus mugo EM-6 144 144 74 LC
Pinus muricata NA-23 49 12 VU
Pinus nelsonii MCW-30 108 108 6 EN
Pinus nigra EM-8 145 24 94 LC
Pinus occidentalis MCW-69 132 8 EN
Pinus oocarpa MCW-7 96 54 LC
Pinus palustris NA-30 52 43 EN
Pinus parviflora MAJ-158 273 34 LC
Pinus patula MCW-15 100 100 24 LC
Pinus peuce EM-12 148 14 NT
Pinus pinaster EM-9 146 37 LC
Pinus pinceana MCW-32 109 13 LC
Pinus pinea EM-10 147 147 30 LC
Pinus ponderosa NA-21 47 48 133 LC
Pinus praetermissa MCW-8 96 8 NT
Pinus pringlei MCW-18 101 17 LC
Pinus pseudostrobus MCW-9 97 97 75 LC
Pinus pumila MAJ-144 264 265 50* LC
Pinus pungens NA-35 54 18 LC
Pinus quadrifolia NA-13 42 7 LC
Pinus radiata NA-19 46 5 EN
Pinus remota MCW-29 107 14 LC
Pinus resinosa NA-7 39 52 LC
Pinus rigida NA-29 52 52 35 LC
Pinus roxburghii MAJ-17 185 19 LC
Pinus rzedowskii MCW-31 109 109 2 VU
Pinus sabiniana NA-24 50 22 LC
Pinus serotina NA-34 54 22 LC
Pinus sibirica MAJ-9 179 37* LC
Pinus squamata MAJ-37 202 1 CR
Pinus strobiformis MCW-26 106 39 LC
Pinus strobus NA-8 40 40 95 LC
Pinus sylvestris EM-5 143 216* LC
Pinus tabuliformis MAJ-42 204 66 LC
Pinus taeda NA-26 50 61 LC
Pinus taiwanensis MAJ-127 253 7 LC
Pinus tecunumanii MCW-12 99 32 VU
Pinus teocote MCW-23 104 48 LC
Pinus thunbergii MAJ-154 271 26 LC
Pinus torreyana NA-18 46 3 CR
Pinus tropicalis MCW-68 132 4 VU
Pinus uncinata EM-7 145 18 LC
Pinus virginiana NA-28 51 51 49 LC
Pinus wallichiana MAJ-18 185 186 36 LC
Pinus wangii MAJ-40 203 2 EN
Pinus ×densithunbergii ‒ ‒ NE
Pinus ×hakkodensis ‒ ‒ NE
Pinus ×neilreichiana ‒ ‒ NE
Pinus ×rhaetica EM-6 144 4 NE
Pinus yunnanensis MAJ-43 205 39 LC
Platycladus orientalis MAJ-107 241 12 NT
508 tables and indices

Table (cont.)
Species Map number Page Photo page Degree squares IUCN Red List
Podocarpus acuminatus SA-28 447 3 NT
Podocarpus acutifolius AU-54 379 6 LC
Podocarpus affinis AU-121 424 4 NT
Podocarpus angustifolius MCW-72 134 5 CR
Podocarpus aracensis SA-28 447 2 LC
Podocarpus archboldii MA-27 307 12 VU
Podocarpus atjehensis MA-29 308 3 NT
Podocarpus borneensis MA-19 304 4 LC
Podocarpus bracteatus MA-12 301 9 LC
Podocarpus brasiliensis SA-24 444 14 LC
Podocarpus brassii MA-23 305 305 26 LC
Podocarpus brevifolius MA-61 328 1 NT
Podocarpus buchii MCW-75 136 6 EN
Podocarpus capuronii AF-20 470 6 EN
Podocarpus celatus SA-16 438 17 LC
Podocarpus chingianus MAJ-84 227 1 DD
Podocarpus confertus MA-21 305 4 EN
Podocarpus coriaceus MCW-73 135 11 LC
Podocarpus costalis MA-14 302 5 EN
Podocarpus costaricensis MCW-61 127 1 CR
Podocarpus crassigemma MA-25 306 15 LC
Podocarpus cunninghamii AU-51 377 41 LC
Podocarpus decumbens AU-105 413 1 CR
Podocarpus deflexus MA-16 303 2 EN
Podocarpus dispermus AU-34 364 2 NT
Podocarpus drouynianus AU-29 361 4 LC
Podocarpus ekmanii MCW-74 135 2 LC
Podocarpus elatus AU-31 362 20 LC
Podocarpus elongatus AF-19 469 11 LC
Podocarpus fasciculus MAJ-175 283 11 VU
Podocarpus gibbsiae MA-62 328 2 VU
Podocarpus glaucus MA-13 302 13 LC
Podocarpus globulus MA-18 304 4 EN
Podocarpus glomeratus SA-10 435 15 LC
Podocarpus gnidioides AU-109 415 2 NT
Podocarpus grayae AU-32 363 363 9 LC
Podocarpus guatemalensis MCW-60 126 31 LC
Podocarpus henkelii AF-19 469 8 EN
Podocarpus hispaniolensis MCW-74 135 136 2 EN
Podocarpus humbertii AF-21 471 3 EN
Podocarpus insularis AU-125 426 12 LC
Podocarpus lambertii SA-26 445 446 24 NT
Podocarpus latifolius AF-19 469 469 25 LC
Podocarpus laubenfelsii MA-20 304 5 EN
Podocarpus lawrencei AU-37 365 365 14 LC
Podocarpus ledermannii MA-24 306 15 LC
Podocarpus levis MA-16 303 9 LC
Podocarpus longifoliolatus AU-107 414 4 EN
Podocarpus lophatus MA-30 308 2 VU
Podocarpus lucienii AU-104 412 5 EN
Podocarpus macrocarpus MA-28 307 6 EN
Podocarpus macrophyllus MAJ-86 228 86 LC
Podocarpus madagascariensis AF24, 25, 26 472 15 NT
Podocarpus magnifolius SA-17 439 16 LC
Podocarpus matudae MCW-58 124 26 VU
Podocarpus micropedunculatus MA-22 305 4 NT
Podocarpus milanjianus AF-18 469 33 60 LC
Podocarpus nakaii MAJ-130 254 4 EN
 tables and indices 509

Table (cont.)
Species Map number Page Photo page Degree squares IUCN Red List
Podocarpus neriifolius AU-124, MA-7426, 298 299 182 LC
Podocarpus nivalis AU-53 378 378 23 LC
Podocarpus novae-caledoniae AU-108 414 414 3 LC
Podocarpus nubigenus SA-35 454 454 19 NT
Podocarpus oleifolius SA-18 439 57 LC
Podocarpus orarius AU-126 427 6 NT
Podocarpus palawanensis MA-28 307 1 CR
Podocarpus pallidus AU-122a+b 425 3 VU
Podocarpus parlatorei SA-11 436 436 14 NT
Podocarpus pendulifolius SA-15 438 3 EN
Podocarpus perrieri AF-23 472 3 CR
Podocarpus pilgeri MA-10 300 55 LC
Podocarpus polyspermus AU-106 413 2 EN
Podocarpus polystachyus MA-9 300 48 VU
Podocarpus pseudobracteatus MA-26 306 9 LC
Podocarpus purdieanus MCW-76 137 1 EN
Podocarpus ramosii MA-29 308 3 DD
Podocarpus ridleyi MA-17 303 5 VU
Podocarpus roraimae SA-19 440 6 LC
Podocarpus rostratus AF-22 471 4 EN
Podocarpus rubens MA-8 299 24 LC
Podocarpus rumphii MA-11 301 51 NT
Podocarpus rusbyi SA-12 436 7 VU
Podocarpus salicifolius SA-14 437 437 7 LC
Podocarpus salignus SA-37 455 456 16 VU
Podocarpus salomoniensis AU-123 425 4 EN
Podocarpus sellowii SA-25 445 14 EN
Podocarpus smithii AU-33 364 2 LC
Podocarpus spathoides MA-30 308 2 DD
Podocarpus spinulosus AU-30 362 362 16 LC
Podocarpus sprucei SA-13 437 8 EN
Podocarpus steyermarkii SA-20 441 7 LC
Podocarpus subtropicalis MAJ-85 228 1 DD
Podocarpus sylvestris AU-103 412 6 LC
Podocarpus tepuiensis SA-21 441 10 LC
Podocarpus teysmannii MA-15 302 20 NT
Podocarpus totara AU-52 377 377 32 LC
Podocarpus transiens SA-27 446 7 EN
Podocarpus trinitensis MCW-77 137 1 NT
Podocarpus urbanii MCW-76 137 1 CR
Prumnopitys andina SA-38 456 9 VU
Prumnopitys exigua SA-8 434 4 NT
Prumnopitys ferruginea AU-55 379 32 LC
Prumnopitys ferruginoides AU-97 407 5 LC
Prumnopitys harmsiana SA-6 433 18 NT
Prumnopitys ladei AU-35 364 364 1 VU
Prumnopitys montana SA-7 434 21 VU
Prumnopitys standleyi MCW-62 127 3 EN
Prumnopitys taxifolia AU-56 379 33 LC
Pseudolarix amabilis MAJ-64 215 10 VU
Pseudotaxus chienii MAJ-122 250 250 11 VU
Pseudotsuga japonica MAJ-165 277 277 3 EN
Pseudotsuga macrocarpa NA-46 60 60 6 NT
Pseudotsuga menziesii NA-45 59 152 LC
Pseudotsuga sinensis MAJ-61 213 214 45 VU
Retrophyllum comptonii AU-101 410 5 LC
Retrophyllum minus AU-102 411 411 1 EN
510 tables and indices

Table (cont.)
Species Map number Page Photo page Degree squares IUCN Red List
Retrophyllum piresii SA-28 447 1 DD
Retrophyllum rospigliosii SA-9 435 13 VU
Retrophyllum vitiense MA-58 325 16 LC
Saxegothaea conspicua SA-36 455 21 NT
Sciadopitys verticillata MAJ-176 284 14 NT
Sequoia sempervirens NA-64 71 71 10 EN
Sequoiadendron giganteum NA-65 72 5 EN
Sundacarpus amarus MA-59 326 327 64 LC
Taiwania cryptomerioides MAJ-89 230 231 11 VU
Taxodium distichum NA-66 72 81 LC
Taxodium mucronatum MCW-56 122 123 32 LC
Taxus baccata EM-42 169 32, 170 155 LC
Taxus brevifolia NA-91 87 60 NT
Taxus canadensis NA-92 88 87 LC
Taxus chinensis MAJ-123 251 37 EN
Taxus contorta MAJ-32 194 20 EN
Taxus cuspidata MAJ-189 291 43 LC
Taxus floridana NA-93 88 1 CR
Taxus globosa MCW-57 123 12 EN
Taxus mairei MAJ-124 251 51 VU
Taxus wallichiana MAJ-33 195 36 EN
Tetraclinis articulata EM-41 168 26 LC
Thuja koraiensis MAJ-149 267 6 VU
Thuja occidentalis NA-69 74 75 78 LC
Thuja plicata NA-69 74 56 LC
Thuja standishii MAJ-180 286 18 NT
Thuja sutchuenensis MAJ-92 233 233 1 EN
Thujopsis dolabrata MAJ-178 285 34 LC
Torreya californica NA-94 89 16 VU
Torreya fargesii MAJ-119 248 22 EN
Torreya grandis MAJ-120 248 249 23 LC
Torreya jackii MAJ-121 249 6 EN
Torreya nucifera MAJ-190 292 34 LC
Torreya taxifolia NA-95 89 1 CR
Tsuga canadensis NA-52 64 75 NT
Tsuga caroliniana NA-53 64 7 NT
Tsuga chinensis MAJ-65 216 216 53 LC
Tsuga diversifolia MAJ-169 280 17 LC
Tsuga dumosa MAJ-23 189 186 44 LC
Tsuga forrestii MAJ-66 217 8 VU
Tsuga heterophylla NA-50 62 83 LC
Tsuga mertensiana NA-51 63 63 56 LC
Tsuga sieboldii MAJ-168 279 16 NT
Widdringtonia cedarbergensis AF-8 463 464 2 CR
Widdringtonia nodiflora AF-7 463 25 LC
Widdringtonia schwarzii AF-9 464 2 VU
Widdringtonia whytei AF-10 464 464 1 CR
Wollemia nobilis AU-11 351 351 1 CR
Xanthocyparis nootkatensis NA-78 80 32 LC
Xanthocyparis vietnamensis MAJ-140 260 261 3 EN
 tables and indices 511

Other Maps

Map title Map number Page

Global distribution of all conifer species GTC-1 11
The distribution of degree cells with ≥ 6 genera per cell GTC-2 13
The distribution of degree cells with ≥ 10 taxa per cell GTC-3 13
The distribution of degree cells with rarity values GTC-4 13
Distribution of conifer species in North America (N of Mexico) and Greenland NA-1 35
The distribution of species in Pinaceae (black), Taxaceae (yellow) and NA-2 36
Cupressaceae (red) in the contiguous United States
Map of North America showing numbers of conifer genera per one degree cell NA-3 37
Map of North America showing numbers of conifer species per one degree cell NA-4 37
Rarity values for 1 degree cells (North America) NA-5 38
The distribution of the genus Picea in North America (N of Mexico) NA-36 55
The distribution of the genus Abies in North America (N of Mexico) NA-54 65
Distribution of the genus Juniperus in North America and Greenland NA-79 80
Distribution of conifer species in Mexico, Central America and the West Indies MCW-1 91
Map of Mexico, Central America and the West Indies showing numbers of conifer MCW-2 92
genera per one degree cell
Map of Mexico, Central America and the West Indies showing numbers of conifer MCW-3 92
species per one degree cell
Rarity values for 1 degree cells (Mexico, Central America and the West Indies) MCW-4 92
The distribution of conifer species in Mexico MCW-5 94
The distribution of species in the genus Pinus in Mexico MCW-6 95
The distribution of species in the genus Abies in Mexico and Central America MCW-34 111
The distribution of species in the genus Juniperus in Mexico and Guatemala MCW-43 116
The distribution of conifer species in Central America MCW-59 126
The distribution of conifer species in the West Indies and Bermuda) MCW-63 129
Distribution of conifer species in Europe and the Mediterranean EM-1 140
Map of Europe and the Mediterranean showing numbers of conifer genera per EM-2 141
one degree cell
Map of Europe and the Mediterranean showing numbers of conifer species per EM-3 141
one degree cell
Rarity values per 1 degree cell (Europe & Mediterranean) EM-4 142
Distribution of conifer species in mainland Asia and Japan MAJ-1 173
Map of Mainland Asia and Japan showing numbers of conifer genera per MAJ-2 174
one degree cell
Map of Mainland Asia and Japan showing numbers of conifer species per MAJ-3 174
one degree cell
Rarity values per one degree cell (Mainland Asia and Japan) MAJ-4 174
The distribution of conifer species in mainland China and Taiwan MAJ-35 199
The distribution of the genus Picea in mainland China and Taiwan MAJ-48 208
The distribution of the genus Abies in mainland China and Taiwan MAJ-72 220
The distribution of the genus Juniperus in mainland China and Taiwan MAJ-97 237
The distribution of conifer species in Taiwan MAJ-125 252
The distribution of conifer species in Japan MAJ-150 268
Map of Japan showing numbers of conifer genera per one degree cell MAJ-151 269
Map of Japan showing numbers of conifer species per one degree cell MAJ-152 270
Rarity values per one degree cell (Japan) MAJ-153 270
Distribution of conifer species in Malesia MA-1 294
Map of Malesia showing numbers of conifer genera per one degree cell MA-2 295
Map of Malesia showing numbers of conifer species per one degree cell MA-3 295
Rarity values per 1 degree cell (Malesia) MA-4 296
The distribution of species of Podocarpaceae (black) and of species belonging MA-5 297
to other families (red) in Malesia
The distribution of the genus Podocarpus in Malesia MA-6 298
Distribution of conifer species in Australasia AU-1 339
Map of Australasia showing numbers of conifer species per 0.5 degree cell AU-2 342
Rarity values per 0.5 degree cell (Australasia) AU-3 342
The distribution of conifer species in Australia AU-4 345
512 tables and indices

Table (cont.)
Map title Map number Page
The distribution of conifer species in New Zealand AU-46 373
Number of species in 1/10 degree cells (New Caledonia) AU-68 388
The distribution of the genus Araucaria in New Caledonia AU-75 392
The distribution of conifers on the ‘interarc’ islands of the SW Pacific AU-115 420
Distribution of conifer species in South America SA-1 430
Map of South America showing numbers of conifer genera per one degree cell SA-2 431
Map of South America showing numbers of conifer species per one degree cell SA-3 431
Rarity values per one degree cell (South America) SA-4 431
The distribution of conifer species in Brazil SA-22 442
The distribution of conifer species in South America south of 30° S SA-29 448
Distribution of conifer species in Africa AF-1 460
Map of Africa showing numbers of conifer genera per one degree cell AF-2 461
Map of frica showing numbers of conifer species per one degree cell AF-3 461
Rarity values per one degree cell (Africa) AF-4 461
The distribution of the genus Podocarpus in Africa AF-17 468
Distribution of dated herbarium collections of Podocarpaceae EFR-1 475
Map of the world with 10 degree graticules showing the global distribution of all EFR-2 478
conifers
Map of the world with TDWG Level-4 units showing IUCN Red List threatened EFR-3 479
taxa (VU, EN, CR) per unit area
Map of the world with the distribution of threatened (pink and red) and non- EFR-4 480
threatened (white) conifer taxa
Map of Sumatera showing current land use and the localities of rainforest conifers EFR-5 482
Conifer diversity of the Sunda Shelf based on 37 stacked SDMs at 5 arc minute EFR-6 484
spatial resolution
Conifer diversity on the Sunda Shelf, aggregated to 1 degree spatial resolution EFR-7 485
The distribution of the species Araucaria montana in New Caledonia plotted EFR-8 488
using Google Maps

List of Tables

Title of table Number Page

Herbaria with >1000 records in the Conifer Database 1 3
Salient features in BRAHMS relevant to the Conifer Atlas project 2 7
Salient features of DIVA GIS relevant to the Conifer Atlas project 3 8
The distribution of 26 genera of conifers occurring in Australasia 4 340
The species of Araucaria in New Caledonia and their substrates 5 398
Statistics of ‘continents’ as defined in the Atlas compared 6 476–477
Number of species in five range classes for the eight ‘continents’ 7 477
Numbers of species in the different IUCN Red List Categories 8 479
The predicted status of conifer species in the rainforests of Sumatera 9 481
Countries with numbers of taxa, endemics and IUCN Red List categories of threat 10 495–499