Early Mesozoic Land Flora of Northern Hemisphere

143
The Early Mesozoic Land Flora of the Northern Hemisphere
Sidney Ash
Department of Geology
We.ber State College
Ogden, Utah 84408
INTRODUCTION
In this chapter the, early Mesozoic land flora of the northern

hemisphere is discussed briefl~ and some of the subtle but significant
evolutionary changes that took place during the time are summarized.
Comments about climatic implications of the fossils are also included as
well as comments about the' environments of deposition of the fossils.
Representative plant megafossils of the Early Mesozoic are illustrated.
The Early Mesozoic as used here includes all of the Triassic and Jurassic
Periods and the Neocomian Stage of the Cretaceous.
The flora of the Early Mesozoic, as used here, is included in what has
been called the Mesophytic flora (Gothan and Weyland, 1954). The
Mesophytic flora is the transitional flora that developed after the older
or Paleophytic flora of the Paleozoic and before the modern Cenophytic
flora. The Paleophytic flora, lasting from the acquisition of the land
habit by plants during the Silurian through the Early Permian, was
dominated by lycopsids, sphenopsids, ferns, and primitive gymnosperms.
The Mesophytic flora which was dominated by advanced gymnosperms,
extended from the Late Permian to about the end of the Early Cretaceous
when the flowering plants became common. The Cenophytic flora continues
to the present and is dominated by the flowering plants.
Although transitional in nature, the Mesophytic flora is significant

and noteworthy not only because of the intrinsic value of the plants it
contains but because the flowering plants evidently originated from some
lineage that was present in it (see Beck, 1976; Crane, 1985; Hughes,
1976; Retallack and Dilcher, 1981). The Mesophytic flora is interesting
and quite distinctive as it includes many new taxa in addition to a
number of holdovers from the Paleophytic flora. These holdovers are more
common in the Triassic floras than in the Jurassic and Early Cretaceous
floras and most seem to have become extinct by Late Cretaceous time.
Generally the late Mesophytic flora is more modern in aspect than the
early Mesophytic flora. Some of the new taxa in the Mesophytic flora
(e.g., some of the seed ferns) are quite strange and difficult to classify
and apparently did not evolve into any living forms. Other new taxa
(some of the ferns, cycads and conifers for example) are obviously close
ancestors of certain modern plants and are easy to classify.
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GEOGRAPHICAL AND STRATIGRAPHICAL DISTRIBUTION
The best generally available report on the distribution of Early

Mesozoic land floras was published over ten years ago (Barnard, 1973).
Earlier Dobruskina (1970) had covered the distribution of Triassic floras
in the northern hemisphere in some detail. More recently she
(Dobruskina, 1980, 1982, 1986) has expanded on her discussion of that
topic. Many short reports on the distribution of Early Mesozoic floras
in parts of the northern hemisphere have appeared. They include reports
on the Early Mesozoic floras of Japan (Oishi, 1940; Kimura, 1980), the
Late Triassic floras of Japan (Kimura, 1986) and North America (Ash,
1980), the Jllrassic floras of much of Europe (Wesley, 1966), Iran and
adjacent areas (Assereto et al.·.,1968) and Russia (Vakhrameev, 1970), the
Early Cretaceous floras of Russia (Vakhrameev, 1966, 1970, 1971) and of
Japan (Kimura, 1975). A comprehensive report on the Triassic floras of
the world is now in preparation by John and Heidi Anderson.
In his 1973 report Barnar~ shows with the aid of four paleogeographic
reconstructions, that the early Mesozoic floras of the northern hemisphere
generally occur from about 20 degrees north latitude to very close to the
paleonorth pole of the time. Generally they are irregularly distributed
throughout that zone but they seem to be most common in Central Asia and
Siberia during each period.
Early and Middle Triassic floras are fairly scarce and those that are
present are small and occur mainly in western Europe (Grauvogel-Stamm,
1978) and Siberia (Mogutscheva, 1973; Dobruskina, 1982). Interestingly,
none are known in North America and adjacent regions at all. Late
Triassic floras are much more widely distributed than those of the Early
and Middle Triassic. They include large floras in both the eastern
United States (Ash, 1980; Bock, 1969; Cornet and Olsen, 1986; Fontaine,
1883; Ward; 1900) and the western United States (Ash, 1970a, 1972b, 1978,
1980; Daugherty, 1941) as well as in Mexico (Weber, 1986a, b, c) and
Greenland (Harris, 1931, 1932a, b, 1935, 1937; Pedersen and Lund, 1980).
In addition there are many Late Triassic floras in western Europe
(Harris, 1931; Dobruskina, 1982; Krausel, 1959), in all parts of Russia
(Dobruskina, 1970, 1982, 1986), in China (Hsu and others, 1979; Kimura,
1985) and in other parts of Asia including Korea (Kawasaki, 1925, 1926;
Kimura, 1986), Japan (Oishi, 1940; Kimura, 1980, 1985), and Viet-Nam
(Zeiller, 1902; Srebrodolskaja, 1969).
Early Jurassic floras are fairly widely distributed, particularly in

Italy (Wesley, 1956, 1958, 1966), Israel (Lorch, 1967), Egypt (Ash,
1972a), Greenland (Harris, 1931b, 1932a, 1932b, 1935, 1937; Pedersen and
Lund, 1980), Scandinavia (Lundblad, 1950, 1959), Poland (Raciborski,
1891, 1892), and Germany (Gothan, 1914; Harris, 1931a,). On the other
hand Middle Jurassic floras are not very common. They are best known
from England (Harris, 1961b, 1964, 1969, 1979; Harris and Miller, 1974;
Harris and Millington, 1974), southern Mexico (Wieland, 1914-16; Person
and Delevoryas, 1982). Several Middle Jurassic floras have been
described from the western United States (Ward, 1900) but are in need of
reevaluation. Late Jurassic floras are more common and are more widely
distributed than Middle Jurassic floras. Some large diverse ones occur
in Central Asia and Siberia (Seward, 1912; Vakhrameev, 1966, 1970). A
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small fragmentary Late Jurassic flora is known from Scotland (Van Der
Burgh and Van Konijnenburg, 1984). In addition several significant Late
Jurassic floras occur in China (Wu and others, 1980, Zhang; 1982).
Surprisingly, only a few small Late Jurassic floras are known from North
America (Bell, 1956; Brown, 1972, 1975) although strata of that age are
fairly widely exposed in the western interior region.
Early Cretaceous floras seem to be comparatively scarce everywhere

except in western Europe (Baten, 1975; Hughes, 1975; Watson, 1969, 1977),
eastern Siberia (Krassilov, 1967; Vakhrameev, 1970, 1971; Vakhrameevand
Doludenko, 1961), Japan (Kimura, 1980), and China. Only a few generally
small Early Cretaceous floras occur in North America (Bel~, 1956;
Fontaine, 1889; Doyle and Hickey, 1976; Lapasha and Miller, 1984, 1985;
Miller and Lapasha, 1984).
PALEOECOLOGICAL IMPLICATIONS
Barnard (1973) concluded that a broad equatorial floristic belt and a

polar floristic region were present throughout most of the early
Mesozoic. The floras in the equatorial floristic belt, which are
characterized by abundant ferns, cycads and cycadeoids, suggest a
tropical to subtropical climate with abundant precipitation. On the
other hand the floras in the polar region have fewer ferns, cycads, and
cycadeoids and the seed ferns, conifers, and ginkgoes dominate,
suggesting a somewhat drier and slightly cooler climate. The bo~ndary
between the two regions seems to have shifted southward about 30 0 during
the Triassic and Jurassic from where it had been at the beginning of the
Mesozoic.
An exception within the polar floristic region may have been in

Siberia during the Early Cretaceous. Here the climate probably was more
humid as indicated by the many ferns, cycads, and cycadeoids that occur
there. Apparently the region served as a refuge for plants that had
become extinct elsewhere (Vakhrameev, 1971). The occurrence of floras in
Siberia and on Ellesmere Island in the Late Triassic and Early Jurassic
from localities that were 60· to 70· north at that time presents the
problem of adapt ion to continuous or nearly continuous darkness during
the winter months.
SYSTEMATIC REVIEW
A review of each of the main groups of land plants present during the
Mesophytic in the northern hemisphere follows. General descriptions of
some of the commonly occurring fossils in the groups are also provided
but the details are necessarily brief. Additional morphological and
anatomical information can be found in recent paleobotanical books (e.g.,
Andrews, 1961; Taylor, 1981; and Stewart, 1983). In this section I
follow the scheme of classification used by Taylor (1981).
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Lycopods (Division Lycopodophytina)
Apparently the lycopods were only minor members of the Mesophytic

flora as their remains are comparatively rare in the Early Mesozoic of
the northern hemisphere. Most of them are the remains of herbaceous
forms that closely resemble living genera. Because of this, some have
been assigned to living genera such as Lycopodium and Se1aginel1a (fig.
la) while others have been placed systematically in allied fossil taxa,
such as Lycopodites, Selaginel1ites, and Isoetites.
Some of the most interesting lycopod fossils in the Mesophytic flora

are the remains of larger forms resembling the giant arborescent lycopods
of the Pennsylvanian. The best known example of this growth habit is
Pleuromeia from the Early and Middle Triassic of Eurasia and Australia
(Retallack, 1975). This plant had a thick stem (about 10--25 cm in
diameter) that was as much as 2 meters tall. A crown of narrow, linear
leaves and a large cone were present at the apex of the stem and leaf
scars from the dehiscence of leaves covered the stem (fig. Ie). Although
it was thought that this plant was a xerophyte which inhabited deserts
(Magdefrau, 1931, 1956) reevaluation of the evidence now indicates that
Pleuromeia usually lived in"mangrove swamps"similar to those occupied by
the Pennsylvanian arborescent lycopods (Krassilov and Zakharov, 1975a,
1975b). However, a stunted form in the Lower Triassic of China is
thought to have lived in a desert oasis (Wang and Wang, 1982).
The occurrence of large lycopodoaceous spores, cones, and stem

impressions at scattered localities in the Late Triassic - Early
Cretaceous of the northern hemisphere indicates that other arborescent
lycopsids besides Pleuromeia existed during the Early Mesozoic
(Retallack, 1975). However, little is known about them and it appears
that arborescent lycopsids were always rare during the Early Mesozoic and
finally became extinct during the Early Cretaceous.
Horsetails (Division Sphenophyta)
Unlike the lycopods the horsetails were somewhat more common in the
early Mesozoic of the northern hemisphere as they consistently occur
tllere in most Mesophytic floras. Generally these plants are most
commonly preserved in overbank deposits in deltaic and estuarian systems.
Many of the Mesophytic horsetails were small herbaceous forms that

closely resemble the living Equisetum and are placed in either that taxon
or Equisetites (fig. Id, If). Other herbaceous forms include Phyllotheca
and Schizoneura (fig. Ib). Although common components of gondwanan
assemblages, neither was ever very common in the northern hemisphere and
both appear to have become extinct, at least in the northern hemisphere,
before the end of the Jurassic (Ash, 1986). They are more common in the
Permian and Triassic of the southern continents suggesting that they
originated there and migrated to the northern continents during the Early
Triassic, if not before.
147
d e
b
Figure 1. Representative early Mesozoic lycopods (a, c) and horsetails

(b, d-f). Redrawn from various sources. a, Selaginella, x5,
b, Schizoneura, xl, c, Pleuromeia, xl/lO, d, Equisetites, x1/2,
e, Neocalamites, x1/4, f, Equisetites, x1/2.
=======================================================================
Arborescent horsetails that are considered to be Paleophytic holdovers
commonly occur in many early Mesophytic floras. Typically they are
referred to the genus Neocalamites (fig. Ie). Stems of this plant which
range up to 40 cm or more in diameter and possibly 6 meters or more in
height OCC\lr commonly in the Late Triassic formations of the southwestern
United States (Ash, 1980). The last well documented occurrence of
Neocalamites seems to be in the Middle Jurassic of Yorkshire (Harris,
1961).
Ferns (Division Pteridophyta)
The status of the ferns at the beginning of the Mesozoic is unclear

but it seems that they were scarce following their precipitous decline in
the Permian. The few that were preserved were members of some of the
groups that had been present in the Late Paleozoic (Arnold, 1964). Later
in the Triassic and Jurassic, several new families aros~ and the ferns
expanded in number and diversity eventually coming to constitute a
significant portion of the Mesophytic land flora. However, they once
again declined in importance in the late part of the Early Cretaceous as
the flowering plants expanded. In general, the Mesophytic ferns have a
148
more modern appearance than the Paleozoic ferns and most can be assigned
to living families without question. Many of the Early Mesozoic ferns
have pinnate leaves in which the pinnu1es are arranged in two ranks along
the sides of the stems as in the probable Late Triassic fern Pekinopteris
(fig. 2i). Some have leaves in which the pinnu1es extend outward in a
palmate fashion from the top of stems as in the common Early Mesozoic
ferns C1athropteris (fig. 2h) and Phlebopteris (fig. 2j).
Marattia1es. This order of Paleozoic ferns was also present in small

numbers during the Early Mesozoic. Triassic representatives of the order
include Danaeopsis and Marattiopsis (fig. 2a). The latter genus and
Marattia also occur in the Jurassic and Early Cretaceous.
Filica1es. The Order Filica1es includes seven living families six of

which were present during the early Mesozoic (Taylor, 1981). The first
of these, the Osmundaceae was also present during the Paleozoic and was
represented in the Mesophytic flora by several genera. Interestingly,
this family which had been small during the Late Paleozoic, became one of
the largest and most diversified of the Mesophytic fern families (Arnold,
1964). Mesozoic members of the family include Todites (fig. 2c), some
species of Cladophlebis and possibly Osmundopsis and Cladotheca. The
Gleicheniaceae, which had evolved during the Late Paleozoic was also
present but insignificant during the Early Mesozoic. Gleichenites (fig.
2f) which occurs at several localities in the Jurassic in the northern
hemisphere (e.g., England), is the principal Mesozoic representative of
this family.
One of the first of the new families of ferns to evolve during the
Mesozoic was the Dipteridaceae. This family first appears in the Middle
Triassic of Australia (Webb, 1982). Thereafter it quickly spread
throughout the world as it is quite common in the Late Triassic of North
America and Early Jurassic of Japan and other areas in the northern
hemisphere. Unlike most of the other Mesophytic ferns the leaves in this
family are palmately compound as in C1athropteris (fig. 2h) and
Phlebopteris (fig. 2j). Another important fern family that arose early
in the Mesozoic is the Matoniaceae (Arnold, 1964). This family is found
for the first time in the Late Triassic in the northern hemisphere where
its members became common during the Early and Middle part of the
Jurassic. Subsequently the family seems to have declined significantly
in n\lmbers and variety. The leaves in this family are also pa1amate1y
compound. The most characteristic and widely distributed matoniaceous
fern found in the Late Triassic and Jurassic is Phlebopteris (fig. 2j).
Other Jurassic members of this family are Matonidum and Selenocarpus and
possibly Piazopteris (fig. 2e).
========================================================================
Figure 2. Representative early Mesozoic ferns. Redrawn from various

sources. a, Marattiopsis, xO.5, b, Hausmannia, xl, c, Todites, xl,
d, Klukia, xO.5, e, Piazopteris, xl, f, Gleichenites, xl,
g, Dictyophyllum, xl, h, Clathropteris, xl, i, Pekinopteris, xO.5,
j, Phlebopteris, xl.
149
a d
150
The Schizaeaceae probably evolved during the early Mesozoic as no

megafossils found in older rocks can be safely assigned to the family
(Taylor, 1981). An early member of this family is Klukia (fig. 2d) which
first appears in the Middle Jurassic of Poland and Yorkshire (Harris,
1961). A rather large and important family of Jurassic ferns is the
Dicksoniaceae which is represented by the common genus Coniopteris and
the rare genera Kylikipteris and Eboracia. The Cyatheaceae appears to
have arisen in the Jurassic as petrified stems and leaf stalks that are
referable to this family are known from that period in India. Specimens
are known from the Early Cretaceous in Japan, but the family was never
very abundant during the early Mesozoic. Cibotiocaulis is a silicified
stem from the Early Cretaceous of Japan that is assigned to the family.
Seed Ferns (Division Pteridospermophyta)
As they are now recognized, the early Mesozoic seed ferns of the
northern continents are a small but rather heterogeneous group of plants
that usually have thick leaves and distinctive reproductive structures.
They were never very common in the northern hemisphere during the early
Mesozoic and few, if any, survived there after the Early Cretaceous.
Although many of the organs of these interesting plants are known it is
not yet possible to reconstruct the whole plant. Some of them may have
been shrubs or small trees. The Mesophytic seed ferns of the northern
hemisphere are assigned to three orders: Peltaspermales, Corystospermales
and Caytoniales. The first of these, the Peltaspermales, was a holdover
from the Paleozoic having been rather common in the Late Permian of the
southern continents (Townrow, 1960) and the northern continents
(Dobruskina, 1975). They were minor members of the Late Permian and Late
Triassic floras of Eurasia. However, the order then disappeared rather
suddenly at the end of the Triassic (Dobruskina, 1975). The bipinnate
leaves of these plants are assigned to Lepidopteris (fig. 3b),
Scytophyllum, Vittaephyllum, and other genera including possibly some
species of Callipteris. The seed bearing organ is Peltaspermum and the
pollen-bearing organ is Antevsia (fig. 3c). Pollen grains produced by
this organ are small monosulcate grains (fig. 3c, right).
The second of these families is the Corystospermales which is common

in the Triassic of the southern continents (Thomas, 1933) and generally
rare in the northern hemisphere (Stewart, 1983). There they seem to be
very common in the Middle Jurassic of Yorkshire where the leaves are
assigned to Pachypteris (fig. 3a) and the pollen-bearing organs are
called Pteruchus.
The Caytoniales is the third family of early Mesozoic seed ferns found
in the northern hemisphere. Members of the family are most common in
Eurasia where they first occur in the latest Triassic (Rhaetian Stage) of
Greenland and Scandinavia (Harris, 1932). Members of the family become
increasingly common in younger strata in Eurasia until they reached their
zenith during the Middle Jurassic. They are particularly common in the
famous Middle Jurassic flora of Yorkshire (Harris, 1964). The family
then declined rapidly and disappeared sometime in the Cretaceous. Some
of the last representatives of the order in the northern hemisphere occur
in the Early Cretaceous (Aptian) rocks of Montana (Lapasha and Miller,
151
1985) and in the Early Cretaceous (Albian) of Japan (Kimura, 1975). The
distinctive palmately compound leaves are assigned to Sagenopteris (fig.
3d), the pollen-bearing organ to Caytonanthus (fig. 3e), and the seed-
bearing organ to Caytonia (fig. 3f). The bisacate pollen produced by the
male organ (fig. 3e, right) is assigned to Vitreisporites or Alisporites.
The seeds are small oval bodies without obvious external features (fig.
3f, right).
===========================~===========================================
~A)
0:0 "
f O&J
Figure 3. Representative early Mesozoic Seed Ferns. Kedrawn from

various sources. a, Pachypteris, xO.5, b, Lepidopteris, xO.5,
c, pollen organ Antevsia, x5, and pollen, x450, d, Sagenopteris, xl,
e, Caytonanthus, x2, and pollen, x300, f, Caytonia, xl, and seeds,
x3.
=======================================================================
Cycads (Division Cycadophyta)
The Mesophytic flora includes a large number of once pinnate and non-
pinnate leaves that were once thought to be fossil representatives of the
living cycads. When it was learned (Nathorst, 1902; Thomas and Bancroft,
1913) that the leaves could be consistently divided into two groups on
the basis of their cuticular structure it was evident that the fossils
actually represented two groups of unrelated plants which are now called
cycads and cycadeoids (sometimes called bennettites). In the stomata of
152
the cycads the guard cells are surrounded by three or more so called
subsidiary cells and the cell walls of the epidermis are typically
straight to slightly wavy (see fig. 4b). In the stomata of the
cycadeoids each of the guard cells is flanked by a single subsidiary cell
and the side walls of the epidermal cells are usually sinuous (see fig.
4e). There are other differences. For example, in the living cycads the
reproductive structures are borne at the apex of the stem whereas in the
majority of cycadeoids they are borne on short branches on the sides of
the stems.
The cycads which apparently evolved during the Late Paleozoic were
scarce in terms of numbers and diversity during the Early Mesozoic in
comparison to the cycadeoids. During the Triassic the cycads were
present in the northern hemisphere in very small numbers. The group
became somewhat more common there during the Jurassic and Early
Cretaceous but then declined rather abruptly. Most of the Early Mesozoic
cycads had pinnate leaves such as the Upper Triassic form Bjuvia and the
Jurassic forms Ctenis, Pseudoctenis (fig. 4c) and Paracycas. Some had
non-pinnate leaves that are called Nilssonia (fig. 4a). Generally cycad
leaves are fairly large. For example some specimens of Nilssonia are at
least one meter long and Ctenis may have been as long, if not longer.
Cycad reproductive structures include Palaeocycas from the Upper Triassic
and Beania and Androstrobus (fig. 4a) from the Middle Jurassic.
Apparently none of the Early Mesozoic cycads had very large trunks. The
oldest known cycad stems occur in the Late Triassic of North America and
show two different growth forms (Ash, 1985). In Charmorgia the stem is
short and squat and probably was no more than 30 em tall whereas in
Lyssoxylon and Leptocycas (fig. 4d) the stems are slender and may have
been a meter or more in height (Ash, 1985). A recent reconstruction of
Leptocycas by Delevoryas and Hope (1971) shows it to have had a crown of
leaves and many persistent leaf bases on the lower parts of the stem
(fig. 4d). Harris (1961a) suggested that the plant which bore the leaf
Nilssonia tenuinervis and the pollen cone Androstrobus wonnacotti had a
slender branched stem (fig. 4a). Many of the cycads in the Mesophytic
flora probably grew along the banks of streams and lakes in deltas and on
floodplains. The presence of cycads in a flora seems to indicate a hot,
moist climate (Vakhrameev, 1971).
=======================================================================
Figure 4. Representative early Mesozoic Cycads (a-d) and Cycadeoids (e-

m). Redrawn from various sources. a, reconstruction of cycad plant
bearing the leaf Nilssonia tenllinervis and the seed bearing structure
Androstrobus wonnacotti from Harris (196Ia). The stem is imaginary in
this reconstruction. b, generalized drawing of a cycad stoma, x500,
c, Pseudoctenis, xl, d, reconstruction of plant bearing the leaf
Leptocycas, e, generalized drawing of a cycadeoid stoma, x500~
f, Wielandiella, xl, g, Williamsonia, xl, h, Dictyozamites, x2,
i, Nilssoniopteris, xl/4, j, Otozamites, xl, k, Zamites, xl/3,
1, Pterophyllum, xl/3, ffi, reconstruction of the cycadeoid plant
Williamsonia sewardiana bearing the leaf Ptilophyllum and the
reproductive structure Williamsonia.
153
~
~ 9
154
Cycadeoids (Division Cycadeoidophyta)
Although this division may have evolved sometime during the Middle
Triassic or earlier, the oldest confirmed remains occur in the Carnian
Stage of the Late Triassic at several places in the northern hemisphere
(Ash, 1980). The order then expanded and diversified and became a
dominant in the land flora of the northern hemisphere during the Middle
Jurassic (Harris, 1973). The most common pinnate leaves in the Late
Triassic and Jurassic are Otozamites (fig. 4j) and Zamites (fig. 4k).
Others are Pterophy1lum (fig. 41), Ptilophyllum and Anomozamites. In the
Early Jurassic the genus Dictyophyllum (fig. 4h) appeared and then became
common in the Middle Jurassic. Nilssoniopteris (fig. 4i) is the name
generally applied to non-pinnate leaves in the Mesophytic flora.
Reproductive structures include Wielandiella (fig. 4f) and Williamsonia
(fig. 4g). They first occur in the Late Triassic and another,
Weltrichia, appears in the Middle Jurassic. The foliage and reproductive
structures are generally scarce in the Late Jurassic and Early Cretaceous
whereas cycadeoid stems, which are uncommon in the Triassic and most of
the Jurassic, are common in the Late Jurassic and Early Cretaceous.
Limited evidence suggests that the majority of the Late Triassic-Middle
Jurassic plants in this order had a slender branching habit as shown in
fig. 4m (Delevoryas and Hope, 1976). Squat fleshy cycadeoid trunks first
appear in the Late Jurassic at a few localities in Utah, Wyoming and New
Mexico and on the Isle of Portland in England where they were once called
"fossil birds nests". Such trunks are particularly common in the Early
Cretaceous of Maryland and many western states as well as in several
European countries (Wieland, 1906, 1916). These in situ occurrences are
found preserved in floodplain and delta deposits mainly but some of the
English ones are in coastal deposits (Francis, 1983). The presence of
cycadeoids in the Jurassic and Early Cretaceous floras of the USSR
suggest a hot, moist climate to Vakhrameev (1971).
Conifers (Division Coniferophyta)
A bewildering number of fossil and living taxa are included in the

conifers. Perhaps because of this, their relationships and the
evolutionary history of the group as a whole are somewhat unclear at this
time (cf. Meyen, 1984; Miller, 1977, 1985; Rothwell, 1985).
Nevertheless, it is evident that the conifers were generally much more
widespread and diverse during the Early Mesozoic than they are now. And
at that time they also dominated some of the plant communities that were
then present (Miller, 1977). During the Early Cretaceous or possibly
during the Late Jurassic, however, they began a decline in both diversity
and numbers and their distribution became more and more restricted.
Many of the Early Mesozoic taxa, especially those from the Jurassic
and younger strata are assigned to living families. Fossil evidence
indicates that the direct ancestors of the living families appeared one
by one beginning in the Early Jurassic or possibly earlier, and by the
Early Cretaceous all the families were represented by either fossil or
extant genera (Miller, 1977). In general aspect the early Mesozoic
conifers were fairly similar to living conifers and they seem to have had
155
similar growth habits ranging from shrubs to large trees. Most of them
also had small scale or needle leaves similar to tbose pf the living and
familiar juniper and pine (see figs. 5a-5h). Both the extinct and living
taxa are mainly differentiated and classified by the morphology and
anatomy of their seed cones which generally have distinctive bract-scale
complexes (e.g., figs. 5j-5w). Many species and genera have been
established on the basis of sterile foliage like that shown in figs. Sa -
5h, and 5x. Since it is usually difficult to determine the relationships
of such fossils they are not discussed here.
Cordaites (Cordaitopsida)
This Class was well represented during the Late Paleozoic,

particularly during the Pennsylvanian and Permian, and some Early
Mesozoic fossils tentatively have been referred to the order (Taylor,
1981). The namesake of the order is the large strap-like leaf called
Cordaites that was borne typically by generally tall trees having
coniferous wood. Some species were restricted to the Late Paleozoic
peat-swamp communities. The same general type of leaves occur in the
Mesophytic flora but they have often been referred to other genera,
especially Pelourdea, because there is some question about their true
relationships. A recent discovery in the Upper Triassic of Utah shows
that the leaf Pelourdea was actually borne on a short, herbaceous stem
(fig. 5i) and that the plant probably was a member of a riverine
community (Ash, in press). Although nothing definite is known about the
systematic relationships of the Pelourdea plant it may well have been
cordaitalean. Certainly this plant was only locally abundant in the
Triassic of the northern hemisphere.
Conifers (Coniferopsida)
Voltziales. -This extinct order is sometimes called the "transition

conifers" because their reproductive structures seem to be intermediate
between those of the Cordaitales and modern conifers (Miller, 1977). The
order includes two families of primitive conifers, the Voltziaceae and
the Cheirolepidiaceae.
The Voltziaceae is of particular interest because it includes forms

that were possibly intermediate between the Lebachiaceae of the Late
Paleozoic and the living conifers (Miller, 1977). Some members of the
family are known only from the Permian, others such as Glyptolepis (fig.
5j) are known from the Permian and Triassic and some occur only in the
early Mesozoic, and are especially common in the Triassic. The latter
include the seed cones Voltzia (fig. 5k), Pseudovoltzia (fig. 51) and
Swedenborgia (fig. 5m). Although the foliage of the plants that bore
these cones is unknown some species of the leaf shoot Podozamites (fig.
5x) have been linked with Swedenborgia by Harris (1935). One of the most
completely known members of the family is Aethophyllum from the Lower
Triassic of France (Grauvogel-Starnm, 1978). That plant has narrow linear
leaves that are sessile on the stem and the seed cone is a lax structure
with the bract-scale complexes arranged in a loose helix around the axis
(fig. 5n).
156
b c
. _'
f 9
@ /'.
..
')
'~tff
• ".J .,/.;,
.
p
o
J kim n
~~~o
StU v w
Figure S. Sterile foliage (a-i, x), seed cone scales (j-n, p-w), and a
cone (0) of certain early Mesozoic conifers. Redrawn from various
sources. a, Pagiophyllum, x2, b, Brachyphy11um, x2, c, E1atoc1adus,
xl, d, Hirmerie11a, x2, e, Frene10psis, x2, f, E1atides, x2,
g, Cupressinoc1adus, x2, h, Stachyotaxus, xl, i, reconstruction of
the Pelourdea plant, xl/20, j, G1ypto1epis, k, Vo1tzia,
1, Pseudovo1tzia, m, Swedenborgia, n, Aethophy11um, 0, Hirmerie11a
cone, xl, p, Hirmeriel1a, q, Stachyotaxus, r, Pa1issya,
s, Araucarites, t, Do1iostrobus, u, Schizo1epis, v, Pa1aeotaxus,
w, Marksea, x, Podozamites, xl.
=======================;=================================================
The Cheirolepidiaceae (formerly called Hirmere11aceae) is one of the

more completely known families of extinct Early Mesozoic conifers. This
family first appeared during the Late Triassic and became a dominant
element in the Jurassic and Early Cretaceous. It suddenly declined in
importance during the Cretaceous and eventually became extinct in the
Early Tertiary (Upchurch and Doyle, 1981). The foliage of this family
includes several taxa including Frene10psis (fig. Se), Pseudofrenelopsis
157
and some species of Hirmeriella (fig. 5d), Pagiophyllum (fig. 5a) and
Brachyphyl1um (fig. 5b). The male cone is C1assostrobus and the seed
cone is Hirmeriel1a (figs. 50, 5p). Hirmeriel1a is considered to have
been a arborescent conifer (Taylor, 1981). The family probably included
plants which lived in a variety of habitats including possible halophytic
'mangroves" and had a variety of growth habits, from shrubs to large trees.
Interestingly, xeromorphic characters seem to be present in most species
that have been studied in detail (Alvin, 1982; Watson, 1982). There is
good evidence from the Late Jurassic of England that an arborescent
species lived there under relatively harsh, semi-arid conditions
alongside a hypersaline gulf (Francis, 1983).
Coniferales. This class includes one extinct and six living families.
All of the living families were present during the Early Cretaceous
having evolved at various times during the Triassic and Jurassic (Miller,
1977). Details of their relationships are unclear because of the absence
of critical fossil evidence.
The Pa1issyaceae is a small, extinct family of conifers that evolved

during the Triassic and disappeared during the Jurassic (Taylor, 1981).
Stachyotaxus is a Late Triassic member of the family that is known from
Sweden, Switzerland and Greenland (Harris, 1935; Florin, 1958). The
linear leaves of this plant are arranged in two ranks along the branches
(fig. 5h) and the seed cone-scales consist of a pair of short stalk-like
megasporophy11s arranged on either side of a central axis (fig. Sq).
Pa1issya is a mainly Jurassic member of the family in which the leaves
are helically arranged on branches and the seed cone scales consist of
ten stalk-like megasporophy1ls arranged like those of Stachyotaxus (fig.
5r).
The Podocarpaceae is now restricted to the southern continents where

most of their fossil remains have also been found. There is limited
evidence of its presence in the northern hemisphere including some
specimens of Elatoc1adus (fig. Sc) that resemble the foliage of the
family. Taylor (1981) indicates that this evidence may mean that the
family was once more cosmopolitan in distribution than at present. Be
tllat as it may, it is apparent the Podocarpaceae was of little importance
in the Early Mesozoic flora of the northern hemisphere.
The Araucariaceae is also restricted to the southern continents at

present but it was once also present in the northern hemisphere, possibly
in considerable numbers. In fact, an unusually large number of fossils
has been referred to the Araucariaceae family although the evidence for
the assignment is often weak (Miller, 1977). Most of the fossils are
merely fragments of petrified wood that have some of the general
characteristics of Araucarian wood and thus have been assigned to the
form genus Araucarioxylon. However, this type of wood has also been
identified in other gymnosperm families such as the Glossopteridaceae.
Reproductive structures and foliage that almost certainly represents the
family is present in Middle Jurassic of England (Harris, 1979) and
perhaps elsewhere in the Triassic (Miller, 1977). The family expanded
somewhat during the rest of the early Mesozoic but later declined in
abundance and became restricted to the southern continents. Fossil
representatives of the family in the Early Mesozoic of the northern
158
hemisphere include the seed cones Araucarites (fig. 58), Damarites, and
Doliostrobus (fig. 5t), several isolated types of bract-scales, and
various leafy shoots such as some species of Pagiophyllum (fig. 5a).
The early record of the Cupressaceae is generally meager although it

may extend back to the Late Triassic in France (Miller, 1977). An
example of a possible early member of the family is Cupressinocladus
(fig. 5g) from the Early-Middle Jurassic of Israel (Chaloner and Lorch,
1960). Several species of the wood called Cupressoxylon from the
Jurassic and Early Cretaceous seem to be correctly assigned to this
family. It is not until the Early Cretaceous that representatives of the
Cupressaceae become at all common and that examples of living members of
the family are recognizable (Miller, 1977).
The geologic record of the Taxodiaceae seems to extend back to the

Late Triassic but fossil representatives do not become common until the
Jurassic (Taylor, 1981). Such fossils include some remarkable petrified
cones from the Jurassic of Argentina called Pararaucaria although they
probably are not closely related to the Araucaria (Stockey, 1977). Some
of the species of foliage and cones referred to Elatites (fig. 5f) from
the Middle Jurassic of England may well belong in this family (Miller,
lq77).
The Pinaceae appears to have been the most recent family of conifers
to evolve. According to Miller (1977) it seems certain that Pinus had
appeared during the Early Cretaceous as Alvin (1960) has correctly
assigned some seed cones from the Early Cretaceous of Belgium to the
genus. The identification of Cedrus alaskensis from the Early Cretaceous
of Canada by Arnold (1953) also seems to be well founded according to
Miller (1977). Some seed cones called Compsostrobus and associated
pollen cones and foliage from the Late Triassic of North Carolina have
been assigned to the family by Delevoryas and Hope (1973). Miller feels
that the situation with regard to the other fossils assigned to the
family is less certain.
The Cephalotaxaceae appears to have arisen during the early Mesozoic

but the evidence is not clear concerning the exact timing of the event
(Miller, 1977). The leafy shoot Thomasiocladus from the Middle Jurassic
of England is probably correctly assigned to the family (Harris, 1979).
However, leafy shoots that have been assigned to the extant genus
Cephalotaxus have been identified from the Upper Triassic of Arizona by
Brown (in Stewart and others, 1972) and in younger strata at other
. localities. In addition some similar Early Cretaceous fossils have been
assigned to Cephalotaxopsis (Miller, 1977).
Taxales. -This small extant order may have arisen during the Triassic but
the oldest fossils that can be confidently attributed to it are some
leafy twigs with attached fruits called Palaeotaxus (fig. 5v) from the
Lower Jurassic of Denmark (Florin, 1958). The order is well represented
in the Middle Triassic by several species some of which can be placed in
the modern genera Taxus and Torreya and others which are placed in fossil
genera. These include foliage and seed cones called Marskea (fig. 5w)
and the sterile foliage Storgaardia (Miller, 1977). Additional fossils,
including petrified wood, that can be safely assigned to the order are
159
found in Upper Jurassic and Lower Cretaceous strata at a few localities

in the northern hemisphere. It is apparent from the fossil record that
the Taxa1es have never been very abundant.
Ginkgoes (Ginkgophyta)
Ginkgoales.- The geologic history of this group goes back to the Early
Permian and fan-shaped leaves that compare very closely with those of the
living genus Ginkgo (fig. 6a) are found in rocks of Late Triassic age and
perhaps in even older ones. Such leaves are known from the latest
Triassic of Ellesmere Island and Greenland (Harris, 1935), the Early
Jurassic of Greenland, the Middle Jurassic of Yorkshire (Harris and
Millington, 1974) the Early Cretaceous of Montana (Lapasha and Miller,
1985), and many other Early Mesozoic localities (Stewart, 1983). Several
types of wedge-shaped leaves that are divided into several segments have
also been assigned to the order. The most common of these are Baiera and
Sphenobaiera (fig. 6b). Both occur in some frequency in rocks of latest
Triassic and Jurassic age in the northern hemisphere. A pollen cone and
seeds that compare with those of the living Ginkgo have been described
from the Middle Jurassic of England (see Harris and Millington, 1974).
Czekanowskiales. -The leaves of this extinct group are finely divided

into narrow segments containing a single vein each. The leaves which are
grouped into small clusters at the end of what is called a short shoot
are referred to the genera Czekanowskia (fig. 5c) and Solenites and the
seed cone is called Leptostrobus (fig. 5d). Members of the order appear
for the first time in the latest Triassic in Greenland and become widely
distributed in the Jurassic and Early Cretaceous in Eurasia (Harris and
Miller, 1974). Throughout their time of existence they seem to have
remained northerly in their distribution and are not found, for example,
in central or southern Europe. The order was most widely distributed
during the Middle Jurassic when it occurred in a band extending from
Yorkshire to eastern Siberia. Thereafter it withdrew from the south and
west so that during the Early Cretaceous the order was found only in
north and east Siberia where it soon became extinct (Vakhrameev, 1970).
Incertae Sedis
The Mesophytic flora contains a number of plant fossils that are

difficult to classify at this time because they are strange and key
information is lacking. The unusual features of some of these plants
amply illustrate the strange character of the Mesophytic flora. One of
the most provocative of these is the Glossopteris-like leaf Mexiglossa
(fig. 6e) from the Middle Jurassic of Mexico (Delevoryas and Person,
1975) and Late Triassic - Early Jurassic of Honduras (Ash, 1981).
Although this fossil closely resembles the characteristic Gondwana leaf
Glossopteris it was not assigned to that genus because reproductive
structures typical of Glossopteris have not been found associated with
the Mexican fossils. Another provocative plant found in the E.arly
Mesozoic is Sanmiguelia from several localities in the Late Triassic of
160
Figure 6. Ginkgoes (a, b), Czekanowsk1ales (c, d), and fossils ~f

uncertain classification (e - g). Redrawn from various sources.
a, Ginkgo, xl, b, Sphenobaiera, x1/2, c, Czekanowskia, x1/4,
d, Leptostrobus, xl, e, Mexiglossa, x1/4, f, Sanmiguelia, xl/10,
g, Dinophyton, x1/4.
the southwestern United States (Brown, 1956; Ash, 1980). The morphology
of the entire plant is fairly well known as several specimens that are in
the position of growth have been described from Colorado (Tidwell, et al.,
1977). It has large, plicate, broadly elliptical leaves that are
helically ~rranged on short narrow stems (fig. 6f). This fossil is of
particular interest because it has been regarded by some authors as an
angiosper~altho\lgh others disagree (e.g., Doyle, 1973). Another
possible pre-Cretaceous angiosperm is Furcula from the latest Triassic in
Greenland (Harris, 1932) and Russia (Hughes, 1976). It is based on
lanceolate leaves that divide in the middle into two segments. Although
the cuticular structure that this plant has occurs in angiosperms it also
occurs in other groups so opinions are divided on the status of this
interesting fossil (Scott, et aI, 1960). The remains of an enigmatic
plant called Dinophyton have been described from many localities in the
Upper Triassic of North America (Ash, 1970b). It includes coniferous-
like foliage and a seed-bearing structure that does not match any other
known seed-bearing structure. All surfaces of the foliage and seed-
161
bearing structure are covered with small sharp hairs (fig. 6g).
Pentoxylon is a Jurassic plant from India that is difficult to classify
even though it is fairly well known (Harris, Bose and Pal, 1985). The
vascular tissue in the stems of this plant is usually divided into five
segments arranged in a ring and the non-pinnate leaves are arranged in a
whorl at the apex of the stem. Because it is so difficult to classify
the Pentoxylon plant is sometimes placed in a distinct group, the
Pentoxyleae. Even less clearly understood is Hermanophyton from the
Upper Jurassic of Utah (Arnold, 1962). The vascular tissue in the stems
of this plant is also divided into segments, a character that is
reminiscent of Pentoxylon. The reproductive structures and foliage of
this plant are not known at this time and its classification is
uncertain.
SUMMARY
A. rich distinctive flora was present during the Early Mesozoic. It

included representatives of nearly all major plant groups that are now
living. Some of the plants such as the arborescent lycopods and
horsetails and a few of the ferns and conifers were holdovers from the
Paleozoic. Most of the plants, however, evolved during the early
Mesozoic although they had their roots, so to speak, in the Paleozoic.
Acknowledgements. I am grateful to Dr. R. A. Gastaldo (Auburn) and Dr.

C. A. Miller, Jr. (University of Montana) for constructively criticizing
an early draft of this chapter. I thank Naomi Hebbert for assisting me
with the-illustrations. Research on some of the material described here
was supported by generous grants from the Earth Sciences Section of the
National Science Foundation.

Early Mesozoic Land Flora of Northern Hemisphere

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143

The Early Mesozoic Land Flora of the Northern Hemisphere

In this chapter the, early Mesozoic land flora of the northern

Although transitional in nature, the Mesophytic flora is significant

GEOGRAPHICAL AND STRATIGRAPHICAL DISTRIBUTION

The best generally available report on the distribution of Early

Early Jurassic floras are fairly widely distributed, particularly in

Early Cretaceous floras seem to be comparatively scarce everywhere

Barnard (1973) concluded that a broad equatorial floristic belt and a

An exception within the polar floristic region may have been in

Lycopods (Division Lycopodophytina)

Apparently the lycopods were only minor members of the Mesophytic

Some of the most interesting lycopod fossils in the Mesophytic flora

The occurrence of large lycopodoaceous spores, cones, and stem

Horsetails (Division Sphenophyta)

Many of the Mesophytic horsetails were small herbaceous forms that

Figure 1. Representative early Mesozoic lycopods (a, c) and horsetails

Ferns (Division Pteridophyta)

The status of the ferns at the beginning of the Mesozoic is unclear

Marattia1es. This order of Paleozoic ferns was also present in small

Filica1es. The Order Filica1es includes seven living families six of

Figure 2. Representative early Mesozoic ferns. Redrawn from various

The Schizaeaceae probably evolved during the early Mesozoic as no

Seed Ferns (Division Pteridospermophyta)

The second of these families is the Corystospermales which is common

Figure 3. Representative early Mesozoic Seed Ferns. Kedrawn from

Figure 4. Representative early Mesozoic Cycads (a-d) and Cycadeoids (e-

Cycadeoids (Division Cycadeoidophyta)

Conifers (Division Coniferophyta)

A bewildering number of fossil and living taxa are included in the

This Class was well represented during the Late Paleozoic,

Voltziales. -This extinct order is sometimes called the "transition

The Voltziaceae is of particular interest because it includes forms

The Cheirolepidiaceae (formerly called Hirmere11aceae) is one of the

The Pa1issyaceae is a small, extinct family of conifers that evolved

The Podocarpaceae is now restricted to the southern continents where

The Araucariaceae is also restricted to the southern continents at

The early record of the Cupressaceae is generally meager although it

The geologic record of the Taxodiaceae seems to extend back to the

The Cephalotaxaceae appears to have arisen during the early Mesozoic

found in Upper Jurassic and Lower Cretaceous strata at a few localities

Czekanowskiales. -The leaves of this extinct group are finely divided

The Mesophytic flora contains a number of plant fossils that are

Figure 6. Ginkgoes (a, b), Czekanowsk1ales (c, d), and fossils ~f

A. rich distinctive flora was present during the Early Mesozoic. It

Acknowledgements. I am grateful to Dr. R. A. Gastaldo (Auburn) and Dr.

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