THE GEOLOGICAL HISTORY OF GYMNOSPERMS

Author(s): EDWARD WILBER BERRY

Source: The Plant World , FEBRUARY 1916, Vol. 19, No. 2 (FEBRUARY 1916), pp. 27-41
Published by: Wiley on behalf of the Ecological Society of America

Stable URL: https://www.jstor.org/stable/43477496

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 THE GEOLOGICAL HISTORY OF GYMNOSPERMS*

EDWARD WILBER BERRY

The Johns Hopkins University , Baltimorei Md.

There are two considerations that I desire to empha

fore attempting to sketch the geological history of gym
These are (1) the vast period of earth history that ha
before any recognizable traces of vascular plants are
the record, and (2) the reasons why they have not yet b
in the earlier Paleozoic periods. One is apt to speak of the
Paleozoic flora as if it were an entity comparable to the recent
flora although the Permian gymnosperms were separated from
their known Devonian ancestors by a far greater interval than
separates the existing flora from pre-glacial floras. A chronology
is then one of the most important factors in any understanding
of the succession of floras that have clothed the earth.
There are a great many ways of estimating geological time,
most of them dependent on the comparisons of past processes
with present processes, and all nothing more than approxima-
tions. It is possible, however, to get an idea of the relative dura-
tions of the different periods that will be reasonably accurate.
Geologists segregate the rocks of the earth's crust, into five sys-
tems - Archeozoic, Proterozoic, Paleozoic, Mesozoic and. Ceno-
zoic. If 5 be taken to represent the duration of the Cenozoic
then the Mesozoic will be represented by 12, the Paleozoic by
27, the Proterozoic by 27 and the Archeozoic by 30. Some
idea of the vastness that these figures represent may be gained
from the statement that the Swiss Alps, Himalayas, Pyrenees,
and Rockies have been elevated and the Colorado Canon eroded
since the dawn of the Cenozoic ; and neither mammals nor angio-
sperms are found as far back as the beginning of the Mesozoic.
* Paper read in the Symposium on Gymnosperms at the meeting of the
Botanical Society of America at the University of California, August 3, 1915.
27

THE PLANT WOBLD, VOL. 19, NO. 2, 1916

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28 EDWARD WILBER BERRY

The first land-plants known occur in th

only midway back in the Paleozoic, and
faunas of any representative character o
Paleozoic.
From the figures of ratios given above it is seen that over half
of geologic time had elapsed before any recognizable animals or
plants are found. This explains the relatively high degree of
organization of the earliest known fossils, both animal and vege-
table, since more time intervenes between the formation of the
earliest recognizable Archeozoic rocks and the appearance of
fossil plants in the known record, than intervenes between that
time and the present. This situation really holds out the hope
that any day we may discover some unmetamorphosed lens of
shale in these older rocks that will reveal to us the life of the
ages long antecedent to the Cambrian.
From the high organization and consequent differentiation of
the earliest known floras and faunas it is a legitimate conclusion
that their ancestors passed through a long antecedent period of
evolution. Terrestrial plant life in the oldest records comprises
representatives of five phyla of vascular plants,1 including
seed-plants, so that the lost interval represents a longer period
of evolutionary activity than the known record down to the
present, thus confirming the evidences of chronology derived
from physical sources.
The bulk of the early sediments have been many times sub-
jected to the action of epeirogenic and orogenic forces and are
consequently so altered that all traces of contained organisms
have been obliterated. Moreover terrestrial plants are almost
invariably preserved in lacustrine, fluviatile, or palustrine de-
posits on the bosoms of the continents or in the estuary and
lagoon deposits of the continental margins. True marine de-
posits of whatever age rarely contain plants except such as con-
tributed to the sea-drift and these are relatively few in number
and usually unrecognizable by the time they were buried.
1 The Pteridophyta or ferns, the Lepidophyta or lepidodendrons and sigil-
larias, the Arthrophyta or calamites and their allies, the Pteridospermophyta or
seed-ferns, and the Coniferophyta or conifers.

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 GEOLOGICAL HISTORY OF GYMNOSPERMS 29

Both the deposits formed on the surfaces of the con

and those formed along their margins are the first to be d
in subsequent cycles of erosion, of which there have bee
Such types of sediments form a relatively small per cent
geological column. The earliest unmistakable deposits
sort known are in the Devonian and these do contain the remains
of a complex terrestrial flora. In the following pages I have
used the term "gymnosperm" in a morphological and not a
taxonomie sense for there are serious objections to including in a
single phylum all those very diverse plants that agree in possess-
ing a micropylar canal.
PALEOZOIC HISTORY

Botanical literature abounds with discussions of the o

mutual relations of the gymnosperms. The older views were
based for the most part on the reproductive organs of the exist-
ing species while the later interpretations lean heavily on vascu-
lar anatomy, and both are highly speculative and have led equally
competent students in quite opposite directions.
We all seem to share to a more or less degree that human ten-
dency to consider only certain structures and not the totality of
structures. Within my recollection the morphology of the higher
plants was simply the morphology of floral parts as it is largely
to this day. The monumental system of Engler and Prantl is
based almost entirely on the morphology of what might properly
be called the accessory reproductive organs. There is a de-
voutly held tradition among botanists that the reproductive
organs are the most conservative of all organs and therefore
merit most consideration. A priori I would consider them as
more subject to evolutionary change than almost any other part
of the plant and much less conservative than, for example, vas-
cular structures or even foliar characters. Geologic history, it
seems to me, fully confirms this assumption. A few examples
will make this clear. Heterospory has arisen independently in
nearly all the great groups. Its logical sequel - the seed habit,
has also been acquired in unrelated lines, e.g., in the great group
of Lepidophytes which culminated in Paleozoic times, Lepido-

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30 EDWARD WILBER BERRY

carpon and Miadesmia are almost as far

habit as the contemporaneous Pteridosperms. Among the
Pteridosperms we have the typical filicalian foliage, both in
form and structure, surviving the change from homospory
through heterospory to seed formation. Among the cycado-
phytes we find the foliage assuming the existing form in the
Carboniferous and continuing unchanged throughout the ages
during which the plants passed through the series of vascular
and floral changes represented by the Mesozoic Cycadeoidales
(Bennettitales) and Williamsoniales and the modern Cycadales.
Are the gymnosperms in the widest sense of the term mono-
phyletic or polyphyletic? It seems to me that this is a question
of degree rather than kind. Obviously if evolution is true we
only have to go back far enough to find common ancestors. In
the sense that none of the forms classed as gymnosperms are
genetically related except in the remotest way to the group in-
cluding the Lycopods and their Paleozoic relatives (which groiip
may properly be called the phylum Lepidophyta) or to the group
including the equiseta and their Paleozoic relatives (which group
may be properly be called the phylum Arthrophyta) the gymno-
sperms are certainly monophyletic.
In another sense they may with propriety be considered poly-
phyletic and for the following reasons: The Paleozoic Pterido-
sperms while they grade off into little known forms of quite un-
suspected structural possibilities, exhibit in those forms that are
better known a sum of characters which precludes them from
forming an order in a single taxonomie group that shall be
termed Gymnospermae, and it seems to me that they are appro-
priately to be considered a separate phylum - the Pteridosper-
mophyta, as suggested by Ward. Their sole coniferous character
is the micropylar avenue of fertilization, a quite necessary stage
in the evolution of the seed habit, as is proved by Lepidocarpon
and Miadesmia among the Lepidophyta. It is true that the
Pteridosperms do clearly indicate the character of the ancestral
stock that leads to existing cycads, but they are probably not
directly related, and in the latter we have another group, enor-
mously diversified during the Mesozoic, one which in the judg-

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 GEOLOGICAL HISTORY OF GYMNOSPERMS 31

ment of many students is also entitled to rank as a ph

under the name of Cycadophyta as suggested by Nath
There remain four groups commonly considered as ord
probably of independent fern ancestry: (1) the Paleozo
daitales, (2) the rather circumscribed Ginkgoales, which r
from the late Paleozoic to the Recent, and enjoyed a co
able adaptive radiation during the Mesozoic, (3) the G
which Lignier would .consider angiosperms, and which W
would apparently derive from the cycad alliance, and whi
be passed over because they have no known fossil repr
tives, and (4) the Coniferales. These four groups, or a
the first two and the last, should be united in a larger
phyletic rank for which the term Coniferophyta prop
Coulter may suffice.
The oldest known floras, those of the middle and later
zoic, contain two very distinct and yet not entirely un
types of gymnosperms - the Pteridospermae or seed fe
the Cordaitales.
The Pteridospermae are the more primitive of the two and
show every evidence of fern ancestry. Their stems were known
for a generation or more before the detached fronds and seeds
were correlated with them, and these stems because of their
synthetic morphological characters were long known as Cycado-
filicales. The Seed Ferns had foliage that was entirely fern-
like in form and habit, as for example in the Sphenopteris type,
which was highly decompound. These fern-like fronds had
always been considered to be those of ferns although Stur as
long ago as 1883 had suggested that they might be cycadaceous.
The seeds were large and complex but were borne on fronds that
were but slightly modified from the ordinary vegetative type
and the pollen was borne in what might well be termed synangia
on similarly unmodified fronds. Throughout the whole group
nothing approaching cones is known, and the anatomy of stem or
leaf, or both, was of the fern type. There was naturally great
diversity of detail in this large plexus of forms and some are
much more fully known than others.
At the present time at least a dozen genera based on stem

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32 EDWARD WILBER BERRY

anatomy and segregated into five famili

Pteridospermae ( Cladoxylon , Volkelia, Ly
gium, Megaloxylon, Calamopitys, Medullosa
Cycadoxylon, Ptychoxylon, Protopitys).
involved in this group with their correspo
as follows:

Sphenopteris - Lagenostoma Callipteris - Carpolithus (in part)

Neuropteris - Rhabdocarpus (in part)C allipteridium - Tripler ospermum
A lethopteris - Pachytesta Doleropteris - Codonospermum
Odontopteris - Odontopterocarpus A neimi tes - W ardi a
Linopteris - Trigonocarpus (in part)Dicksonites - Carpolithus (in part)

According to Scott, the Pteridosperms exhibit two lines of

evolution, - one toward polystely, which becomes elaborated
(Medullosaceae) and then extinct; and another in which a single
stele becomes elaborated (Lyginodendraceae), the latter type is
considered as approaching the Cycadophytes. According to
Chodat, Lyginodendron is a specialized fern, an end product not
concerned with cycad ancestry, which group originated from the
Medullosaceae (which he renames Protocycadaceae).
We know too few members of what was really a great group
to be dogmatic, but there can be no question but that our whole
conception of Paleozoic floras and ancestral seed plants has been
profoundly altered by what has thus far been demonstrated.
The other great group of Paleozoic gymnosperms, exempli-
fied by the genus Cordaites, while they also show in their structure
the evidences of a fern ancestry, had travelled the evolutionary
road much farther than the Pteridosperms. They had lost all of
the more obvious fern characters and were tall trees with a thick
cylinder of secondary wood ( Dadoxylon ) of the araucarian type,
differing from modern conifers merely in their larger pith. Their
foliage consisted of a dense crown of spirally arranged, invari-
ably simple and elongated, and sometimes very large leaves.
The fructifications were small catkin-like cones, the ď1 and 9
separate as in modern conifers. The cones were small, with
thick spirally bracteate axes, in the one case bearing micro-
sporophylls, and except for the bracts comparable to Ginkgo ;
and in the other case bearing short-stalked ovules. The pollen

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 GEOLOGICAL HISTORY OF GYMNOSPERMS 33

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34 EDWARD WILBER BERRY

Fig. 2. Diagram showing the filiation and geologic

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 GEOLOGICAL HISTORY OF GYMNOSPERMS 35

grains continued to grow after leaving the microspo

showed a group of prothallial cells or antheridium whi
oped motile sperms directly and without the form
pollen tube. The seeds were bilaterally symmetrical and
winged.
Although we know Cordaites in detail, the impressions of
foliage indicate that it constituted a large and diversified genus.
Other types are known from fragments showing stem anatomy
{Pity s , Poroxylon, Mesoxylon, Callixylon, Caenoxylon, Meso-
pitys, Parapitys, etc.) and it is quite possible that the Ginkgo
line diverged from somewhere in the Cordaitean plexus in the
late Paleozoic.
Both the Pteridosperm and the Cordaites stocks are known
from the Devonian: Both were dominant groups that became
cosmopolitan in the Carboniferous : Both continue in the Permian
and neither is certainly known from the Mesozoic although a
few stragglers may have lingered on into that era.
Three additional types of gymnosperms appear in the late
Paleozoic record but as our knowledge of both is based largely
upon impressions of the foliage little is known of their exact
affinities. These three groups are the Cycadophyta, the Arau-
cariaceae and the Ginkgoales, all of which became greatly dif-
ferentiated and common during the Mesozoic. The genus
Psygmophyllum of the Devonian Carboniferous and Permian is
supposed to represent the Ginkgoales as is also the genus Whittle-
seya of the Carboniferous. In the Permian a considerable num-
ber of Ginkgo-like forms are known ( Ginkgophyllum , Baiera,
Saportaea, etc.). The araucariaceae are represented in the
Upper Carboniferous by Walchia and Schizodendron and in the
Permian by Walchia, Ulmannia and Gomphostrobus .2 The
cycadophytes appear to be represented as early as the Carbo-
niferous by fronds of Plagiozamites and Pterophyllum, and the
genus Sphenozamites is added in the Permian, but all of these
types are infrequent until the next era.

2 The genus Voltzia of the Permian is often considered as representing the

Taxodieae.

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36 EDWARD WILBER BERRY

MESOZOIC HISTORY

The Mesozoic era has been called the age of conifers, and no
without reason. The gymnosperms are represented during
time by five great groups - the Williamsoniales, Cycadales
Cycadeoidales (Bennettitales) of the cycadophyte phylum,
by the Ginkgoales and Coniferales of the Coniferophyte phy
Of these five groups the Williamsoniales and Cycadeoidal
appear to be limited to the Mesozoic: the Ginkgoales beca
practically extinct during the succeeding era: and there ar
large variety of peculiar Mesozoic types of Coniferales.

Cycadophyta

It is but a few years since the Mesozoic cycadophytes, so

abundantly represented by foliage in all parts of the world from
the Arctic to the Antarctic, were generally accepted as represent-
ing true coniferous and carpellary-leaf bearing cycads not far
removed from existing types. Then came the discoveries of
Carruthers, Solms-Laubach, and especially Wieland, demonstra-
ting the existence of forms in vegetative habit and appearance
wholly cycadean, but with fructifications unique among gymno-
sperms. The sporophylls, instead of being solitary and terminal,
were borne in abundance on lateral dwarfed branches among
the leaf -bases, and were bisporangiate, suggesting comparisons
with Ranalian flowers such as those of Liriodendron or Magnolia.
These forms are properly referable to the genus Cycadeoidea
although many students abroad prefer the later name of Bennet-
tites. These spectacular discoveries have resulted in a vast
amount of misapprehension among botanists, who assume that
the bulk of the Mesozoic cycadophytes were of this type, whereas
it now seems clear that the Cycadeoidales were simply a bizarre
side-line, stereotyped early in the Mesozoic, and without issue.
Their great importance rests in the fact that because of their
silicification at a number of localities they have furnished the
key to the interpretation of the much more abundant and plastic
group - the Williamsoniales. In these latter are represented a
vast plexus of forms showing great variations in size, foliage,

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 GEOLOGICAL HISTORY OF GYMNOSPERMS 37

branching, and sporophyll structure. Their stems were

vailingly slender. Their fronds, of which innumerable g
and species have been described from all parts of the worl
among others the following types: Ptilophyllum, Zami
Otozamites, Sphenozamites, Glossozamites, Sewardia, Dic
mites, some Podozamites, Wielandiella, Nilsonia, Pteroph
Dioonites, etc.
The fructifications, which were the basis for the establish
of the genus Williamsonia long before their botanical af
was known, were borne on an extended bracteate peduncle
which is interpreted as a continuation of the slender stems but
which may sometimes have been a lateral branch. These
sporophylls consist of an ovulate receptacle much like that of
Cycadeoidea surrounded by a whorl of microsporophylls that
are much more reduced than in Cycadeoidea. These form a
fairly well defined reduction series illustrated by the genera
Cycadocephalus, Weltrichia, Williamsonia, and Wielandiella.
Some are unisporangiate and are thought to be whorls from the
cones of either monoecious or dioecious forms. For the details
of the structure and relationships of these forms the reader must
be referred to the recent literature on the subject.
The evidence for the existence of Mesozoic Cycadales of the
modern type is decisive enough, but since the evidence of the fossil
fronds has been invalidated by the discovery that they may rep-
resent any of the cycadophyte orders it seems probable that the
Cycadales themselves were never an extensive group. The evi-
dence for their existence in the Mesozoic rests on fossil carpellary
leaves of the Cycas type in the lower Jurassic and on Zamia-like
cones in the upper Triassic.

Coniferophyta

The Ginkgoales were abundant during the Mesozoic, but the

differentiations were mainly of specific rank and very few varia-
tions of even generic rank occur. The two main types known -
Ginkgo and Baiera - were much alike in fructifications and dif-
fered chiefly in foliage, the former with long petioles and the

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38 EDWARD WILBER BERRY

latter with short petioles and dichotom

lamina, with narrow linear segments. B
during the Mesozoic and Baiera appears
into the Cenozoic although Ginkgo was ve
regions during the latter era.
Of the three families of Coniferales (A
and Pinaceae) the Araucariaceae present
features both anatomically and in sporo
phology than the other two. The fossil m
fied wood, pithrcasts, impressions of twi
scales and seeds. Several Paleozoic gener
mentioned. In the Mesozoic this family
mon all over the world and included forms almost identical with
the existing Araucaria and Dammara as well as a variety of other
types. Thus along with the single seeded ligulate forms are
others with a pair of seeds to the scale {Pseudoaraucaria) , and
others that were three seeded {Protodammara). Araucaria has
been recorded in the Mesozoic from Spitzbergen on the north to
Louis Phillipe Land on the south; from numerous localities in
North America, Europe and Asia; and from scattered localities
in South America, Africa and Australia. The most abundant
forms in the Upper Cretaceous of the Atlantic Coastal plain
from New Jersey to Alabama are leafy twigs and cone-scales
of the Araucaria Bidwelli type, and Dammara cone-scales are
found from Greenland to Texas, and in Europe as well. A con-
siderable number of genera of Araucariaceae are represented in
the Mesozoic of which the best known are perhaps Pagiophyllum
and Geinitzia.
The family Taxaceae was also abundant during the Mesozoic,
both sub-families being represented. Some of the representative
genera are: Nageiopsis of the Lower Cretaceous; Palissya of the
Triassic and Jurassic ; Stachytaxus of the late Triassic which had
large cupuliferous seeds comparable with those of the modern
Dacrydium; Cephalotaxopsis, Protophyllocladus, Palaeotaxus, Tu-
mion, V esquia, etc. The determinations of these forms rests
not alone upon impressions of the foliage but upon seeds and
cones and structural material of all classes, so there can be no

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 GEOLOGICAL HISTORY OP GYMNOSPERMS 39

question but that the Taxaceae were more differentiat

more abundant during the Mesozoic than at the present t
or that in those early days genera were cosmopolitan tha
since become isolated, of which Tumion affords an exc
illustration.
Of the three subfamilies into which the Pinaceae are commonly
segregated both the Taxiodieae and Cupressineae appear to pre-
cede the Abietineae in the geologic record and to be much more
common during the Mesozoic. Sequoia, which survives in such
grandeur in California, is represented continuously from the late
Jurassic onward.
The Actinostrobinae, easily recognized, are represented by
Widdringtonia and Frenela at a majority of the Mesozoic out-
crops. The subfamily Abietineae as compared with thfe other
two subfamilies is relatively modern. There can be no question
of the evidence of the geologic record on this point, for while the
Abietineae are well represented toward the close of the Mesozoic
they are greatly outnumbered and completely overshadowed in
differentiation by the Taxaceae and Araucariaceae and by the
subfamilies Taxodieae and Cupressineae. The record is com-
pletely in accord with the older views that the Abietineae are
the most specialized and most modern conifers, and although
this relationship is questioned by Jeffrey, his theory of the origin
of the Coniferales from a primitive Abietinean stock cannot be
said to have convinced many students who have a first hand
knowledge of paleobotany.
CENOZOIC HISTORY

The Cenozoic era was a time during which the floral ty

essentially modern in character. Their most marked
were their arborescent richness as compared with exist
tions and the very general cosmopolitanism of the d
types. The development of arid conditions over larg
North America, Asia, Africa and South America, and th
effect of the subsequent Pleistocene glaciation, were
factors in the gradual segregation of these Cenozoic flo
more or less detached regions. No traces of Cycadeo

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40 EDWARD WILBER BERRY

Williamsoniales are known from the Cen

are seldom found. Zamites occurs in the Lower Eocene of
Louisiana and in the Oligocene of France. The genus Encep-
halartos has been recorded from the late Oligocene of Greec
and Dion and Cycadites have been somewhat doubtfully dete
mined from the Miocene of southern Europe.
The Ginkgoales are well represented during the earlier Tertiary
I have collected hundreds of beautiful fossil Ginkgo leaves from
regions in western Montana now given over to Spruce and
Cottonwood. Several undoubted species occur over an area
that includes the Isle of Mull, Greenland, Sachalin, Siberia,
Canada as far north as the mouth of the Mackenzie, Wyoming,
Montana, etc. The later Tertiary records available do not con-
tain Ginkgo except one or two instances from the Pacific coast
of the United States and from southern Europe.
All three existing families of Coniferales were represented
throughout the Cenozoic. The Araucariaceae show a progres-
sive shrinking of range toward the south, although Araucaria is
present in the Eocene of Europe and Dammara in Eurasia. The
subfossil records show that the range of both of these genera is
still shrinking at the present time.
Among the Taxaceae, Podocarpus is present in some force in
the European Tertiary, Cephalotaxus foliage and fruits are
present in both Europe and North America. Taxus and to a
less extent Tumion are quite generally distributed. Among the
Pinaceae, Pinus , Glyptostrobus , Taxodium , Arthrotaxus and
Cryptomeria are quite generally distributed in the North Tem-
perate Zone while the genus Sequoia is truly cosmopolitan.
Sequoia is found throughout both the northern and southern
hemispheres and its wood showing perfectly characteristic
structure demonstrates its presence in Europe until the dawn of
Pleistocene glaciation. The accompanying sketch map (fig. 1)
shows the Cenozoic occurrences of this interesting genus.
Taxodium and Glyptostrobus had an equally wide range in the
northern hemisphere but appear to be absent in the southern.
The genera Frenela and Widdringtonia , lumped with Callitris by
Eichler, are not known from North America after the Mesozoic
but they are common forms in the Cenozoic of Europe.

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 GEOLOGICAL HISTORY OF GYMNOSPERMS 41

RECENT HISTORY

The existing Coniferales are prevailingly temperate t

form vast forest areas especially in the north. They
somewhat less southern massing, the two regions being
by a broad tropical belt united only in the Andean a
Indian regions, but continuous in Tertiary times.
It is beyond the province of this paper to discuss the
distribution of gymnosperms but it may be pointed out
distribution is fully explained by the details of geolog
The present occurrences of such genera as Tumion, Lib
and Callitris are readily understandable if their former
tion is taken into the account. The use of the terms endemic
and monotypie becomes then singularly inappropriate when
applied to genera like Taxodium, Sequoia, Cunninghamia,
Cryptomeria, etc.
The accompanying diagram (fig. 2) furnishes a graphic sum-
mary of the foregoing discussion, illustrating not only the filia-
tion of the principal types of gymnosperms, but also their relative
abundance during the different eras.

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