BULLETIN OF

THE BRITISH MUSEUM

(NATURAL HISTORY)

GEOLOGY
VOL ii
1953-1956

PRINTED BY ORDER OF THE TRUSTEES OF
THE BRITISH MUSEUM LONDON : 1953-1956

DATES OF PUBLICATION OF THE PARTS

No. i. June IQ53

No. 2. November ...... 1953

No. 3. November ...... 1953

No. 4. March IQ54

No. 5. September ...... 1954

No. 6. January ...... 1955

No. 7. September ...... 1955

No. 8. February 1956

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CONTENTS

GEOLOGY. VOLUME 2

Page
No. i. The Deer of the Weybourn Crag and Forest Bed of Norfolk. A.

AZZAROLI ........... i

No. 2. A Coniferous Petrified Forest in Patagonia. M. G. CALDER . . 97
No. 3. The Solution of the Piltdown Problem, j. s. WEINER, K. p. OAKLEY &

W. E. LE GROS CLARK ......... 139

No. 4. Some Upper Cretaceous and Eocene Fruits from Egypt. M. E. j.

CHANDLER .......... 147

No. 5. The Carboniferous Flora of Peru. w. j. JONGMANS .... 189
No. 6. Further Contributions to the Solution of the Piltdown Problem.

J. S. WEINER, W. E. LE GROS CLARK, K. P. OAKLEY and others . . 225

No. 7. The Schizaeaceae of the South of England in Early Tertiary Times.

M. E. J. CHANDLER ......... 29!

No. 8. The Brachyopid Labyrinthodonts. D. M. s. WATSON . . . 315

7 JUL 1953

THE DEER OF

THE WEYBOURN CRAG

AND FOREST BED

OF NORFOLK

A. AZZAROLI

BULLETIN OF
THE BRITISH MUSEUM (NATURAL HISTORY)

GEOLOGY Vol. 2 No. i

LONDON : 1953

BULLETIN OF
THE BRITISH MUSEUM (NATURAL HISTORY)

GEOLOGY

The following papers appeared in Volume I (1949-52) :

Price
No. i (1949). The Pterobranch Rhabdopleura in the English Eocene.

H. D. Thomas & A. G. Davis 75. 6d.

No. 2 (1949). A Reconsideration of the Galley Hill Skeleton. K. P.

Oakley & M. F. Ashley Montagu 55.

No. 3 (1950). The Vertebrate Faunas of the Lower Old Red Sandstone

of the Welsh Borders. E. I. White.

Pteraspis leathensis White a Dittonian Zone-Fossil. E. I.

White . . . 7s. 6d.

No. 4 (1950)- A New Tithonian Ammonoid Fauna from Kurdistan,

Northern Iraq. L. F. Spath ....... ios.

No. 5 (1951). Cretaceous and Eocene Peduncles of the Cirripede Euscal-

pellum. T. H. Withers ........ 55.

No. 6 (1951). Some Jurassic and Cretaceous Crabs (Prosoponidae) .

T. H. Withers 5s.

No. 7 (1952). A New Trochiliscus (Charophyta) from the Downtonian

of Podolia. W. N. Croft . .' IOs.

No. 8 (1952). Cretaceous and Tertiary Foraminifera from the Middle

East. T. F. Grimsdale ........ ios.

No. 9 (1952). Australian Arthrodires. E. I. White .... 155.

No. 10 (1952). Cyclopygid Trilobites from Girvan. W. F. Whittard . 6s.

THE DEER OF THE WEYBOURN CRAG
AND FOREST BED OF NORFOLK

BY

AUGUSTO AZZAROLI

(University of Florence)

Pp. 1-96; 50 Text-figs.

BULLETIN OF
THE BRITISH MUSEUM (NATURAL HISTORY)

GEOLOGY Vol. 2 No. i

LONDON: 1953

THE BULLETIN OF THE BRITISH MUSEUM
(NATURAL HISTORY) instituted in 1949, is
issued in five series corresponding to the Departments
of the Museum, and an Historical Series.

Parts appear at irregular intervals as they become
ready. Volumes will contain about three or four
hundred pages, and will not necessarily be completed
within one calendar year.

This paper is Vol. 2, No. i' of the Geological series.

PRINTED BY ORDER OF THE TRUSTEES OF

THE BRITISH MUSEUM
Issued June, 1953. Price Twenty-five Shillings.

THE DEER OF THE WEYBOURN CRAG
AND FOREST BED OF NORFOLK

By A. AZZAROLI

(University of Florence)

SYNOPSIS

The systematics of the Cervidae and the stratigraphy of the fossiliferous Pleistocene deposits
of the Norfolk coast are clarified by a revision of material mainly in the A. C. Savin collection,
in the British Museum (Natural History), and the John Gunn collection, in the Castle Museum,
Norwich. Three successive faunas are distinguished in the Forest Bed Series. The nomen-
clature and relationships of various other fossil deer are also considered.

CONTENTS

Page

INTRODUCTION . .......... 4

GEOLOGY ............ 5

THE PHYLOGENY OF THE CERVIDAE ....... 7

DEVELOPMENT OF THE ANTLERS . . . . . . . .11

SYSTEMATIC DESCRIPTIONS ......... 15

Genus Libralces Azzaroli . . . . . . .15

Libralces gallicus Azzaroli . . . . . . .16

Libralces reynoldsi n. sp. ....... 19

Libralces minor n. sp. ........ 27

Libralces latifrons (Johnson) . . . . . .27

Libralces incertae sedis ....... 30

Genus Capreolus Hamilton Smith ...... 30

Capreolus capreolus (L.) ....... 32

Genus Cervus L. ......... 32

Cervus cf. elaphus L. . . . . . . . -35

Genus Euctenoceros Trouessart . . . . . . -37

Euctenoceros tetraceros (Dawkins) ...... 38

Euctenoceros ctenoides (Nesti) ...... 38

Euctenoceros sedgwicki (Falconer) . . . . . .40

Euctenoceros incertae sedis ....... 45

Genus Megaceros (Owen) ....... 46

The group of Megaceros giganteus ...... 48

The group of Megaceros verticornis . . . . .51

Megaceros verticornis (Dawkins) ...... 53

Megaceros dawkinsi (Newton) . . . . . .61

Megaceros savini (Dawkins) ....... 67

Megaceros, incertae sedis ....... 72

Cervidae incertae sedis . . . . . . . -73

" Cervus " obscurus n. sp. ....... 73

Cervid cf. Dama nestii nestii Forsyth Major .... 79

Cervid cf. Dama clactoniana Falc. . . . . . .81

Species incorrectly recorded from the Forest Bed . .83

CHRONOLOGY ........... 84

SUMMARY . . . . . . . . . . . .90

REFERENCES ........... 91

INTRODUCTION

A REVISION of the Forest Bed deer has been needed for a long time. The descrip-
tions of the most interesting specimens are scattered in the older literature :
Falconer (1868), Dawkins (1887), Newton (1882), Gunn (1891), Harmer (1899).
A few specimens were described more recently by Reynolds (1929-34), but these
works refer mainly to the antlers, and no comprehensive study has been published.
Moreover the series in the British Museum (Natural History) has been considerably
enriched through the careful and long-continued work of A. C. Savin (1861-1948)
of Cromer ; his first collection was acquired by the Geological Department in
1897, and his second in 1945.

The material examined consists of several hundred specimens, mostly in the
Savin collection. I have further studied the important collection made by John
Gunn (1801 — 1890), now in the Castle Museum at Norwich. There are in addition
a few fossils of various origins in the two museums mentioned, in the Museum of
the Geological Survey, in the Natural History Museum at Ipswich and in the private
collection of Mr. J. E. Sainty of West Runton. I have not seen the material of
other private collections, described by Dawkins (1887).

The fossils are highly mineralized, hard, and generally have a peculiar dark brown
colour and a shiny surface; some of them, belonging to the older faunal elements
(see later), are more ferruginous. Some are much worn, but for the most part
they have a well-preserved surface. However, they consist almost exclusively of
isolated bones. Only the skull of Megaceros verticornis, described as Cervus bel-
grandi by Harmer (1899), was found in connection with the first two vertebrae.
This specimen is also exceptional in that the upper portions of its antlers are present.
In the other specimens only the more resistant lower parts have been preserved.
Most of them consist of lower portions of antlers, frontals, horizontal rami of man-
dibles, isolated teeth, limb bones and vertebrae. Brain cases are not uncommon,
but no facial portions of skulls have been preserved, and maxillae are rare. Humeri
and scapulae are represented only by the lower portions, except a heavy humerus
of Libralces reynoldsi.

In the text I have used the abbreviations B.M. (G.D.) for the British Museum
(Natural History), Department of Geology; G.S. for the Museum of the Geological
Survey of England. The registered numbers of the specimens, when quoted in
the explanations of the figures have not been repeated in the text. British Museum
numbers preceded by the name Savin refer to his second collection.

A cknowledgments

I am greatly indebted to Dr. A. T. Hopwood of the British Museum for having
suggested to me the subject of this study and for his invaluable assistance in my

DEER OF WEYBOURN CRAG AND FOREST BED OF NORFOLK 5

work. I wish to thank also Mr. W. N. Edwards, Keeper of the Geological Depart-
ment of the British Museum ; Miss G. V. Barnard, formerly Curator of the Norwich
Museum; Dr. C. J. Stubblefield, Chief Palaeontologist of the Geological Survey,
and Mr. H. E. P. Spencer, Curator of the Museum at Ipswich, for having kindly
given me facilities for studying their collections; Mr. J. E. Sainty of West Runton,
who led me in a field trip on the Norfolk coast; and Dr. K. P. Oakley (London),
Prof. W. B. R. King (Cambridge), Prof. F. E. Zeuner (London), Prof. I. M. Van der
Vlerk (Leiden) and Dr. R. Lagaaij (Leiden) for their criticism of the chronological
section of this work.

GEOLOGY

The Weybourn Crag and Forest Bed Series of the Norfolk Coast

The Weybourn Crag and Forest Bed series, described in Reid's classical memoir
(1890), is exposed at the foot of the cliffs along the Norfolk coast for some miles
east and west of Cromer, and the Forest Bed is exposed also over a short distance
around Pakefield, near Lowest oft. The cliffs consist of incompetent rocks — marine
and freshwater gravels, sands and clays overlain by glacial drift, the whole resting
on a levelled surface of chalk, and erosion is extremely rapid. The exposures con-
tinually change, and Reid's work is an invaluable record of many years of survey,
at a time when the lesser development of coast defences made observation easier.
Study of the stratigraphy is made difficult by rapid lateral changes, by breaks in
the sequence, and by the discontinuity of outcrops. In many places the contorted
glacial drift has disturbed this series, or cut it down to the chalk.

The marine and freshwater series, to summarize Reid's data, rests on the chalk.
At the junction the surface of the chalk is covered by a " stone band " of large
unworn or little-worn flints; this stone band represents no stratigraphical horizon.
In the country around Cromer it is overlain by the " Weybourn Crag," a false-
bedded shelly sand alternating with laminated clay. This crag generally occurs
in patches on the stone band, beneath high tide-mark, between Weybourn and
Trimingham. It " has been traced continuously from Cromer to Weybourn,
except for short distances, where it is cut out by channels of the estuarine Forest
Bed, or has been ploughed out by glacial action, so that the Boulder Clay rests
immediately upon the Chalk" (Reid, 1890: 139). In a section at East Runton
it yielded a rich fauna of molluscs and some mammal bones. In isolated outcrops
its distinction from the Forest Bed is sometimes difficult, so that many authors
overlooked its occurrence. Towards the south-east the Weybourn Crag becomes
more clayey and probably passes laterally into the Chillesford Clay, which in this
section of the coast has been found only in borings, at Mundesley and possibly at
Happisburgh.

The Forest Bed series covers unconformably the Weybourn Crag and has always
a more or less gravelly base. Probably there is a land surface between these two
formations, " for in one place the Crag has a rather weathered appearance; but of
this one cannot be certain" (Reid, 1890: 149).

The Forest Bed series is in three divisions: a " Lower Freshwater Bed," composed
of carbonaceous clays, silt and loam; an "Estuarine Bed" or Forest Bed sensu

6 DEER OF WEYBOURN CRAG AND FOREST BED OF NORFOLK

stricto, composed of false-bedded carbonaceous sands and laminated clays with
mammal bones and drifted tree-stumps; and an " Upper Freshwater Bed," com-
posed of peaty clays and sands. The discontinuous Lower Freshwater Bed is very
seldom preserved, and its observation to-day is still more difficult than at the time
when Reid saw it on the shore north west of Cromer, where it cut through the
Weybourn Crag (1890: 159), and at Trimingham (1890: 163). The Estuarine Bed
is practically continuous. The Upper Freshwater Bed is somewhat variable in
facies and its identification is not always easy. It also is discontinuous and is
separated from the estuarine division by an eroded and deeply weathered surface.

The Occurrence of Mammals

Many of the mammalian remains were found loose on the beach, especially after
storms. Some, however, were actually extracted from the rock in situ. As a rule
they were said to come from the Forest Bed, sensu lato, which is still considered
the chief mammal-bearing horizon; but in the course of the present work it has
become apparent that they are of different ages, so that their occurrence needs
more careful discrimination. Most of the fossils I have examined were obtained
by A. C. Savin of Cromer, who collected for more than fifty years and gave accurate
indications in his catalogue. The specimens I have determined are said to come
from the Forest Bed at W. Runton, E. Runton, Cromer, Overstrand, Sidestrand,
Trimingham, Mundesley, Bacton, and Pakefield; from the beach at Palling and
from the Upper Freshwater Bed at West Runton. A single specimen, a fragment
of a lower jaw of Megaceros dawkinsi, is said to come from the Weybourn Beds at
Weybourn. The specimens from other collections were all stated to come from
the Forest Bed, partly from the same localities as Savin's, partly from other locali-
ties, such as Ostend, Happisburgh, the Walcot gap, and Hopton. However, not
seldom there is only the indication " Forest Bed; Norfolk." A few specimens
were dredged off the coast and on the Dogger Bank.

It is remarkable that Savin did not distinguish an Upper Freshwater Bed in
many localities where it has been recorded by Reid; possibly he was misled by
differences in facies from the corresponding bed at West Runton, and as a matter
of fact the distinction of these horizons in the field is not always easy. Savin dis-
tinguished a Lower Freshwater Bed at East Runton, and Weybourn Beds at East
Runton and West Runton, but recorded from them only fish remains. Here too
he was probably misled by the common belief that mammal remains occurred
only in the Forest Bed and Upper Freshwater Bed. But, as stated above, Reid
found mammal bones also in the Weybourn Crag, at East Runton. Finally, Savin's
attribution to the Weybourn Crag of a lower jaw from Weybourn might be based
merely on the locality and might be equally incorrect.

The mixed character of the " Forest Bed Fauna " led me to suppose that its
members came from different horizons. The characters of the fossils, consisting of
large and well-preserved antlers of deer, rules out any possibility that they were
washed in.

The distribution of the species I have studied shows that there is no sharp break

DEER OF WEYBOURN CRAG AND FOREST BED OF NORFOLK 7

between the fauna of the estuarine Forest Bed and that of the Upper Freshwater
Bed, which, according to my correlations, belong to the great Mindel-Riss inter-
glacial (see later, section " Chronology "), whereas an older fauna may be sharply
distinguished. There is no doubt that it came from an older horizon, separated
from the estuarine bed by a remarkable gap. The widespread occurrence of this
fauna, as well as the high ferruginous content of most of its specimens, rule out
the Lower Freshwater Bed, whilst the Weybourn Crag corresponds to what may be
expected to be their matrix. In fact, most of Savin's specimens are said to come
from the gravel pans and sands on the shore, at some distance from the cliffs; and
Mr. Sainty of W. Runton has in his private collection a beautiful lower jaw of
Libralces gallicus, a typical representative of this earlier fauna, which he extracted
from a patch of shelly crag directly overlying the stone band, on the shore between
West Runton and East Runton, below high tide-mark. This crag, still occurring
in patches but variously exposed owing to the rapidity of erosion, corresponds to
Reid's description of the Weybourn Crag.

Boyd Dawkins, who certainly was not influenced by prejudices about the strati-
graphy, referred a skull of the same species from the Savin collection (1887, pi. i,
fig. i) to the " Weybourn Beds (Forest Bed Series), East Runton, Cromer." Savin's
catalogue gives for this specimen the sole indication " East Runton Forest Bed."

These older species were found only in the western section of the coast, between
Sidestrand and West Runton, where the Weybourn beds are exposed. Very few
specimens came from Mundesley, where the Chillesford Clays are concealed beneath
the beach.

THE PHYLOGENY OF THE CERVIDAE

The early history of the deer is little known. Probably they achieved the greater
part of their evolution in Asia and North America, where the record is fragmentary
and very scanty, during the Pliocene. In Europe their history is largely one of
successive waves of immigration and replacement. Their classification is therefore
vague. Simpson (1945) did not give an exact picture of the phyletic relationships:
for instance, his Odocoileinae are not a phyletic unit, neither are his Cervinae except
in so far as living species are concerned.

In practice only restricted groupings of related genera can be established; the
precise rank of these higher taxonomic units is largely a matter of taste. They
may be called without prejudice subfamilies. Among the living deer seven sub-
families are recognized: Cervinae, Muntiacinae, Odocoileinae, Capreolinae, Rangi-
ferinae, Alcinae, Hydropotinae. As yet hardly any other taxonomic categories
within or above these subfamilies can be established, and if we take into account
also the fossil forms, things become more difficult.

The present work is concerned only with the Cervinae, Capreolinae and Alcinae.

The Capreolinae and Alcinae

The history of the Capreolinae is quite obscure. Some Miocene forms have been
tentatively referred to this group, but they are poorly known (Azzaroli, 1948:

8 DEER OF WEYBOURN CRAG AND FOREST BED OF NORFOLK

46 ff.), and, so far as I am aware, there is no record of this subfamily during the
Pliocene. The dental characters vaguely suggest some affinity with the Alcinae.
The Alcinae are represented in the Quaternary by three genera, Alces, Cervalces
and Libmlces, closely similar in the characters of the dentition and limb bones, but
widely divergent in the skull. They are practically unknown in the Pliocene, but
were already differentiated in the Pontian, where they are represented by Alces
maeoticus.

The Cervinae

The problem of the Cervinae is less simple, and requires more explanation. In
1948 I stated that they were derived from the Pontian Pliocervinae of S.E. Europe
and China; the definition of the Pliocervinae was based on Zdansky (1925, 1927).
A study of the works of Alexejew and Khomenko on the Pontian deer of S.E. Europe
necessitates some modification of this statement.

(a) The Pontian deer from S.E. Europe

In 1913 Alexejew named Procervus variabilis from Pontian deposits near Petro-
vierovka, S. Russia. A more complete description followed in 1915. This species
is distinctly smaller than a fallow deer; its teeth are brachyodont, the upper molars
have a discontinuous cingulum, and the lower molars a well-developed Palaeo-
meryx-iold. Varying with the individual, P4 is either primitive or advanced; the
upper premolars are primitive and the upper canines relatively strong. The skulls
are badly damaged but seem to be of a primitive, rather unspecialized type. The
brain cases are long, the supraoccipital crests weak. The pedicles are set obliquely
and form two ridges at the sides of the forehead. The antlers, of various ages,
are short, stout and much flattened; their form is irregular and extremely variable.
They consist of a flattened, very strong brow tine, sometimes bifurcated at the
top; and a short beam, gradually expanding upwards into a narrow palmation,
ending in two to six small tines. The surface is deeply grooved. The first bi-
furcation takes place near the burr and eventually a small accessory tine is given
off from it. The antlers of younger individuals are forked, and in the first stage
they consist of simple prickets. The limb bones are primitive. The shaft of the
ulna is strong, the upper epiphyses of the radius and tibia are small, and the fore
limb has complete lateral metacarpals, with a well-developed articulation for the
corresponding toes.

In 1914 Khomenko published a description of some Pontian deer from Taraklia,
Bessarabia. His material was scanty and fragmentary, nevertheless he founded
on them three new genera and species, Cervavitus tarakliensis, Cervocerus novo-
rossiae, Damacerus bessarabiae, based on antler characters. Khomenko's specimens
also included some teeth; they showed slight differences which he supposed to be
specific characters. On these three genera Khomenko founded his new subfamily
Pliocervinae.

Khomenko's genera have been variously accepted. Zdansky (1925) supposed
that Cervocerus novorossiae and Damacerus bessarabiae were synonymous; Simion-
escu & Dobrescu (1941) accepted the specific distinction only, and determined other

DEER OF WEYBOURN CRAG AND FOREST BED OF NORFOLK 9

specimens from Bessarabia as Cervocerus bessarabiae. Simpson followed these
authors and recognized the genera Cervavitus and Cervocerus, and included Damacerus
in the latter.

However, there is little doubt that Khomenko's genera and species are all synony-
mous. The differences between the antlers are due to growth stages, and the teeth
do not differ by more than individual characters. The choice for the generic and
specific names is thus open, and here we shall adopt Damacerus bessarabiae. Its
holotype is the antler figured by Khomenko (1914, pi. 4, fig. 6), the only one of
Khomenko's specimens that is fully grown.

The deer from Petrovierovka is closely related to this. The teeth are indis-
tinguishable; P4 displays the same fluctuations. The f rentals, too, seem to be
very similar, and the only marked differences lie in the antlers. In Damacerus
bessarabiae they are less flattened, the brow tine is smaller and the first bifurcation
is set high above the burr. It is consistent therefore to attribute these two species
to the same genus. Procervus is preoccupied and the valid name is Damacerus,
which will include therefore two species, D. bessarabiae (the genotype), and D.
variabilis.

(b) The Pontian deer from China

In 1925 and 1927 Zdansky identified with Khomenko's Cervocerus novorossiae
some deer of various Pontian localities in China. Other specimens from Shansi
were referred to the same species by Teilhard & Trassaert (1937). These identi-
fications are incorrect. The Chinese deer show only superficial similarities to those
of S.E. Europe, but at the same time they display differences showing that they
belong to completely different lineages.

The Chinese so-called Cervocerus novorossiae, as defined by Teilhard & Trassaert
[Zdansky 's description is sometimes vague and partly contradicted by his figures;
moreover some of his attributions are questionable (see Teilhard & Trassaert, p. 38)],
is a species of slightly larger size than the European Damacerus. Its skull is rather
primitive; the pedicles form two prominent ridges on the sides of the forehead.
The lacrimal pits are large and deep. The antlers are three-tined, cylindrical, and
display a great variability in the position of the first bifurcation and in the length
of the beam. Teilhard & Trassaert distinguished a " short type," with a straight
beam, and a " long type," with a gently curved beam; they are linked by inter-
mediate forms. The dentition is brachyodont. P4 is primitive and there is no
trace of the Palaeomeryx-iold on the lower molars. The upper premolars have the
inner wall split, and a strong cingulum is stated to occur in the upper molars, but
this is not to be seen in the figure. There are two complete but very thin lateral
metacarpals, with rudimentary epiphyses.

With the possible exception of some more flattened antlers (pi. 5, figs, i, 2),
Zdansky's Cervocerus novorossiae corresponds on the whole with this species. Ac-
cording to him the cingulum is variable in the upper molars, and in the lower molars
he described what he regarded as the remnants of a Palaeomeryx-i old : " Es handelt
sich meistens um eine, vorn mehr oder weniger scharf begrenzte Rinne. An nur

io DEER OF WEYBOURN CRAG AND FOREST BED OF NORFOLK

ganz wenigen Zahnen ist eine solche nicht zu konstatieren." This is somewhat
vague, but there is certainly no typical Palaeomeryx-iold, and no trace of it can
be seen in the published photographs.

Zdansky also included in the Pliocervinae the genus Procapreolus Schlosser, the
validity of which has been questioned by Teilhard & Trassaert. The latter authors,
on the other hand, described a Cervavitus demissus whose generic identity with the
European forms is no less questionable.

(c) The relationships of the Pontian Cervinae

In conclusion, in the Pontian, apart from some smaller forms more or less closely
related to the Muntiacinae, and from the Alcinae, the Cervidae were already repre-
sented by two well-differentiated groups, namely the European Damacerus, with
two species, and the Chinese three-tined deer, seemingly belonging to one species,
for which new generic and specific names are required. They differed in the charac-
ters of the antlers, of the dentition, and of the fore limbs. These characters enable
us to make some partly hypothetical inferences about their relationships.

In its dental characters the European Damacerus is closely related to the two-
tined Miocene deer, and may have been derived from a genus with a well developed
burr, e.g., Euprox (for full references on Miocene deer see Stehlin, 1939, with biblio-
graphy).

In my opinion the value of the Palaeomeryx-iold. has been underestimated. It is
supposed to have gradually disappeared by the end of the Miocene, but there is no
evidence for this assumption. Throughout the Miocene it does not show any ten-
dency to become reduced or to fluctuate, and it is still uncertain whether it has
really vanished, or has been wiped out by the extinction of the species that bore it.
A solution to this question might possibly be afforded by Cervus ramosus Croizet &
Jobert (1826-28, fasc. 5, pi. 5, fig. 2). The age of the type is uncertain, but the
same species occurs in the Villafranchian of St. Vallier-sur- Rhone and of Villaroya ;
there is no trace of the Palaeomeryx-iold in its lower molars. A more primitive
variety, Cervus ramosus var. pyrenaicus, which might perhaps be the ancestor of the
typical form, was described by Deperet (1897 : 99-112, 124-125) from the Upper
Pliocene of the Roussillon. Among the many lower jaws which can be attributed
to this variety, there are specimens with a distinct Palaeomeryx-iold and specimens
without any trace of it. I am unable to state whether there are intermediate
conditions, nor whether all these jaws belong to the same species ; this however is
highly probable, because the other deer from the Roussillon (Deperet's Capreolus
australis and Capreolus ruscinensis, but actually belonging to one species) are much
smaller. If so, Cervus ramosus would afford a good example of the Palaeomeryx-iold
disappearing by fluctuation.

In the three-tined Pontian deer from China there was no Palaeomeryx-iold. P4
was more primitive than in the European deer of the same age; the upper pre-
molars were rather molarized, but this character always seems to be very variable
and its significance is not quite clear.

This is all that is known of the Upper Miocene ancestors of the Cervinae. A

DEER OF WEYBOURN CRAG AND FOREST BED OF NORFOLK n

large part of them very probably originated from the Chinese species, and the
lineage represented in the Pontian of S.E. Europe may possibly have survived
with Cervus ramosus up to the Lower Pleistocene.

(d) The Pliocene and Pleistocene Cervinae

The history of the Cervinae during the Pliocene is obscure. Besides Cervus
ramosus and its small relative from Roussillon, which might be related to Damacerus,
several species of quite uncertain affinities are known. Most of them were found
in the Montagne de Perrier and other localities of France. There is no evidence
that any of them survived into the Pleistocene.

It would be consistent to give each lineage a distinct generic name, but the use
of the name Cervus, although improper, cannot be dispensed with as long as these
species are so poorly known. However, the name Anoglochis, originally proposed
by Bravard, Croizet & Jobert, might perhaps be adopted for Cervus ramosus.

At the beginning of the Quaternary we find that the various genera which compose
the subfamily are already differentiated, although in many cases represented by
relatively primitive species. Euctenoceros and Megaceros make their first appearance
in the Red Crag of East Anglia, Euctenoceros also in the Dutch Poederlian (see
later), Dama in the Villafranchian of Tuscany (Azzaroli, 1948), Rucervus in the
Pin j or stage of the Siwalik hills (to be described in a forthcoming paper by the
author), Rusa in the Sanmenian of Nihowan (Teilhard & Piveteau, 1930), Sika in
Chouk'outien and other Pleistocene localities (Young, 1932), Axis in the Lower
Pleistocene of Shansi and Java (Teilhard & Trassaert, 1937; Stremme, 1911);
Cervus s.str. makes its first appearance in the second interglacial of Europe. Ela-
phurus is doubtful in the Sanmenian of Nihowan, but has been identified in the
Pleistocene of Japan (Matsumoto, 1915), and is represented also by a subfossil
species at Anyang, N. Honan (Teilhard & Young, 1936). Przewalskium alone is
not known as a fossil. There are in addition some primitive and poorly known
species which cannot be fitted into any of these genera: " Cervus " rhenanus from
Tegelen, " Cervus " perolensis from the Auvergne (Bout & Azzaroli, 1953), " Cervus "
punjabensis and another species from the Upper Siwaliks (to be described in a
forthcoming paper), and " Cervus " philisi from Seneze (Schaub, 1942).

Many of these genera and species may have been derived from the Pontian three-
tined deer of China; but some highly divergent genera, as, e.g., Elaphurus and
Megaceros, are very probably of a quite different origin.

THE DEVELOPMENT OF THE ANTLERS

(a) General Considerations

Much has been written on the growth of the antlers, but our knowledge on this
point is not entirely satisfactory. Each species and genus has its own peculiar
pattern of antlers, obviously determined genetically, but the mechanism by which
this pattern is brought into existence is practically unknown. Wislocki and others
(1946, 1947) have studied the innervation and the process of ossification. It has

12 DEER OF WEYBOURN CRAG AND FOREST BED OF NORFOLK

been established that the form of the antlers is not influenced by the course of
nerve fibres, and attempts to explain the form of the antlers by the course of blood-
vessels (see literature in Wezel, 1949) are equally unsatisfactory. Moreover the
antlers display an individual variability of form and size that cannot be explained
by the action of genetic factors alone. There is evidence that more factors, partly
genetic, partly purely mechanical, influence the determination of their form.
Huxley (1932) established that their size is controlled by a complicated allometric
law, varying from species to species. It is known, too, that increase in size of the
antlers, within a species or subspecies, is followed by a more complicated ramifica-
tion. Inasmuch as this phenomenon occurs among individuals of the same genetic
constitution, or even in successive growth stages of the same individual, it cannot
be because of genetic differences, but rather because of purely mechanical factors :
the antler tissue, growing from the top of the pedicle, seems to be unable to develop
indefinitely in cross-section, but tends to divide as soon as it has reached a certain
limit of bulk. A closer study of this phenomenon is obviously outside the province
of palaeontology.

In the recent deer, division generally takes place dichotomously, with the first
bifurcation or bifurcations in a nearly longitudinal plane. This led Pocock (1933)
to formulate his theory of dichotomous growth; but it may be shown that this
law is not general.

Each species obviously has its own range of variation, and eventually its parti-
cular type of asymmetry, as, e.g., the reindeer. Broadly, the species with more
complicated antlers are also the more variable. Very much has been written on
the variability of antlers, especially of the red deer ; one of the most recent accounts
has been given by Wezel (1949). Allowance must however be made for the un-
natural conditions under which red deer very often live (inbreeding, artificial selec-
tion, unfavourable or exceedingly favourable environment). In deer living under
natural conditions the range of variation is not so wide.

(b) History

If we trace back the history of the deer we may form a mental picture of the
primitive conditions of the antlers, and of the way the more advanced features
gradually became established.

No continuous phyletic line of an appreciable length has yet been ascertained.
The principle of parallel evolution, recently exposed and discussed by Merla (1949 :
117 ff.), will be extensively applied here. This implies the more general principle
of evolution controlled by internal factors, the validity of which has been ques-
tioned by many recent authors who hold a purely " Darwinian " point of view.
The reasons for the interpretation accepted here have been discussed at length by
Merla (1949) and by Watson (1949).

In my opinion the principle of evolution directed by internal factors is the only
one to give a satisfactory explanation of the phyletic development of antlers. It
could be argued that their development might be controlled by factors linked with
characters under selective control ; this interpretation however gives no explanation

DEER OF WEYBOURN CRAG AND FOREST BED OF NORFOLK 13

for the numerous instances of parallel evolution, nor for the polymorphism of antlers,
which is in striking contrast with the uniformity of the other body characters.

Apart from the evolution of antlers, there are not many clear examples of ortho-
evolution of characters which are surely not under selective control. A spectacular
exception is the pachyostosis of the skull of Megaceros.

Lower and Middle Miocene: the primitive antlers

True Cervidae, shedding their antlers, made their first appearance in Europe
during the Burdigalian and remained very primitive during the Helvetian and
Tortonian (for full reference see Stehlin, 1939). They form a polymorphic group.
Complete and well-preserved skulls are not known, but great differences are dis-
played in the position of the pedicles, and in many cases there is no typical burr.
But one common feature is apparent : the antlers never grow in the form of prickets.
Even in the earliest stages they tend to divide, taking the form of an irregular crown
(Stephanocemas ; in fact the form is somewhat intermediate between a crown and
a fork) , a fan (Palaeoplatyceros] or a longitudinal fork (Heteroprox, Dicroceros, Euprox) .
In Asia some of these forms survived into the Pliocene, and may perhaps have given
origin to the living Muntiacinae.

Pontian. The emergence of modern types

A trend towards the formation of a beam became apparent in the Pontian.
Amphiprox, from Eppelsheim, was still very primitive, with short antlers consisting
of a straight beam and a very small brow tine, branching off high above the burr;
but at the same time, deer with more complicated antlers lived in the S.E. of Europe
and in China. They all had a distinct burr and a branched beam.

Damacerus is probably closely related to the earlier European deer. Damacerus
variabilis seems to represent the most primitive condition. The differentiation
between brow tine and beam is not very well established : they are both flattened,
the brow tine is relatively large and bifurcated at the top. The general form of
the antlers is very variable and irregular. The antlers of young individuals consist
of prickets, but it is questionable whether the ontogenetic development actually
reproduces the phyletic history. The formation of the pricket might be due to the
action of genetic factors, normally leading to the formation of the beam, which in
the youngster are not counterbalanced by a sufficiently strong tendency towards
ramification, owing to the small size.

Damacerus bessarabiae is more advanced. There is a typical beam and a much
smaller brow tine. The first bifurcation is set rather high above the burr, and in
my opinion this should be considered an advanced character. The point, however,
is not quite clear, and palaeontological evidence of the evolution of later forms
(see, e.g., Dama nestii nestii from Olivola and the Upper Valdarno; Azzaroli, 1948)
may seem to contradict this assumption. As a matter of fact, genetic and mech-
anical factors trending in opposite directions interact. The upwards shift of the
first bifurcation is determined genetically, and the downwards shift during onto-
genetic development is certainly due to a purely mechanical factor, the larger bulk

i4 DEER OF WEYBOURN CRAG AND FOREST BED OF NORFOLK

of growing tissue inducing earlier bifurcation. This factor also may eventually
act in phyletic evolution. Moreover, a third character, namely, the opening of the
angle of the bifurcations, seemingly under genetic control, may also interfere;
this, however, seems to have become felt only after the end of the Miocene.

In the three-tined Pontian deer of China the antlers consisted of a cylindrical
beam and cylindrical tines. They are rather variable, both in length and in the
position of the first bifurcation. It has been stated in the preceding section that
some palmated antlers figured by Zdansky might perhaps belong to another species.

Pliocene and Quaternary. New evolutionary trends

Although the antlers of these primitive deer were relatively small and simple,
they were very variable. No attempt will be made here to explain this, but in
later forms the patterns of antlers became more constant. New features appeared,
as bifurcations at an obtuse angle, helicoidal tortion and undulation of the beam
and tines, peculiar types of spatial arrangement of the branches (e.g., on a spherical
surface in Rucervus, on a plane in Euctenoceros) , or branching of the tines.

The flattening of the beam and tines is a very common feature. In some cases
it may be a secondary appearance, but its phyletic origin is not always clear. This
is true also of the palmation. In some instances, e.g., Alces dices, no sharp dis-
tinction can be traced between palmated and non-palmated antlers.

(c) On the Homologies of the Tines

The discussion of the formation of antlers leads to the question of the homologies
of the tines. Some words on this problem are necessary because too rigid an inter-
pretation of the homologies would be misleading.

After considering the work of other authors, Pocock (1933) proposed an inter-
pretation of the homologies based on the theory of dichotomous growth. This can
be summarized as follows. The primitive condition is assumed to be represented
by the pricket. In the next stage it divides into an anterior and a posterior tine
(a1 and p1}. These two tines may have the same potentiality of growth and further
division: this happens, e.g., in Elaphurus and Blastocerus. But generally p1 is
the stronger and divides into a2 and pz; then p2 divides into a? and p3, etc. In
accordance with this principle Pocock established the homologies between the
tines of all the living genera of the Cervidae and many fossil forms. In the particular
instance of the red deer the bez tine normally occurs but is not constant; this was
interpreted as having originated from the division of the brow tine.

In 1948 I fully accepted Pocock's views, but I now realize that rigid application
of them is not possible. This theory however has the merit of having shown that
there is no fundamental difference between beam and tines.

It has been shown above that the pricket is not the primitive condition, and that
in the earliest stages branching is more or less irregular. As a matter of fact at
the beginning the tines have no morphological individuality, that is, they cannot
be compared to an organ such as a tooth. What is inherited is the general pattern
of antlers. As this pattern gradually evolves, the beam and the tines acquire a

DEER OF WEYBOURN CRAG AND FOREST BED OF NORFOLK 15

certain degree of individuality, which, however, is not absolutely fixed. The evi-
dence for this is that the eventual suppression of a tine, or the presence of accessory
tines or even an accessory beam (a not uncommon occurrence in the red deer and
reindeer), does not alter the fundamental pattern of the antler. The example of
the tamin (Fig. i) is also instructive. In this species, the surface of the antlers is
sometimes very scabby; the tines (except of course the brow tine) are small, and
no sharp distinction can be traced between the smaller tines and the larger asperities
of the surface.

FIG. i. Rucervus eldi. B.M. (Zool. Dept.), no locality. £ natural size.

True homologies can be established only between related forms, in which the
evolution of the antlers has followed the same path. It is meaningless to argue
whether the homologue of the brow tine of, say, a fallow deer is represented by the
small inner tine of Odocoileus, or by its bifurcated anterior tine, which in turn is
obviously homologous with the anterior tine of Blastocerus ; or to look for the homo-
logue of the bez tine of the red deer in the reindeer and in the giant deer.

SYSTEMATIC DESCRIPTIONS

Genus LIBRALCES Azzaroli

In another paper (1952) I have established the new genus Libralces, with genotype
L. gallicus from the Upper Villafranchian of Seneze. This species is present also
in the Lower Quaternary of the Norfolk coast, together with three other species
of the same genus. These are more imperfectly represented, and are distinguished
mainly by their size. The distinction of the dentitions is easy, but greater diffi-
culties arise in determining other remains, for the most part imperfect fragments

16 DEER OF WEYBOURN CRAG AND FOREST BED OF NORFOLK

of skulls and antlers. The attribution of some of them will therefore remain un-
certain. The limb bones that can be attributed to this genus are scanty.

I have denned the genus Libralces as follows (1952 : 134) : skull broad, depressed
and heavily built; forehead broad; antlers palmated, with a long beam directed
horizontally outwards and gently curved. Nasals long, articulated with the
premaxillae. Upper canines presumably present in the genotype, molars and
premolars brachyodont, lower molars with strong basal columns; traces of the
Palaeomeryx-iold in Mt, eventually also in M2; P4 as in Alces. Forelimb telemeta-
carpal, size large.

• Libralces gallicus Azzaroli
(Figs. 2, 3, 4 B, 5 A, 9 B, 10 D.)

1887. Alces latifrons (Johnson) Dawkins (pars), p. i, pi. i, figs, i, 3 ?, 5.
1931. Alces latifrons (Johnson) : Roman & Dareste, p. 1256.
1944. Alces latifrons (Johnson) : Schaub, p. 285.
1952. Libralces gallicus Azzaroli, p. 134.

Summary Description of the Holotype and Paratype

The type of this species, a complete skeleton of a full-grown male, and a second
incomplete skeleton were found in the Upper Villafranchian of Seneze. The original
description may be summarized as follows: A species of large size, but distinctly
smaller than the living elks. The skull is broad and depressed, with a broad fore-
head and a very thick roof. The face is proportionately less developed than in
Alces and is characterized by the long nasals, which are articulated with the pre-
maxillae. The lower molars bear well-developed basal columns, and the first lower
molar has a distinct groove on the hinder surface of its antero-external crescent.
The antlers consist of a long, slender beam, directed horizontally outwards and
gently twisted, which ends in a small palmation, set obliquely to the axis of the
body, with small tines on its edge like Alces alces. The limb bones are slender and
differ from those of Alces only by their smaller size. The neck is relatively long.

The Specimens from the Norfolk Coast

OCCURRENCE. All the specimens that can be identified with certainty were found
at East Runton and Sidestrand. Imperfect specimens, whose identification is
somewhat doubtful, were found at West Runton, Cromer and Pakefield; the last
are much rolled.

DESCRIPTION. The best specimen is represented by a brain case with its left antler,
originally figured by Dawkins (1887, pi. i, fig. i). In Azzaroli (1952, pi. 15, fig. 2)
and in Fig. 2 the antler has been omitted. An adult, or nearly adult antler from
Sidestrand (Dawkins, 1887, pi. i, fig. 5) and two antlers from East Runton can
also be attributed to this species. All these specimens show a rather narrow range
of variation. The more robust antlers (Fig. 5A, and Dawkins' fig. i) tend to develop
shorter beams.

A second brain case (Fig. 3), mentioned also by Dawkins, very probably belongs

DEER OF WEYBOURN CRAG AND FOREST BED OF NORFOLK 17

FIG. 2. Libralces gallicus, skull from East Runton. B.M. (G.D.), M. 6101. £ natural
size. (Specimen figured by Dawkins, 1887, pi. i, fig. i, and by Azzaroli, 1952, pi. 15,
fig. 2.) A, occipital view ; B, lateral view ; c, basal view.

FIG. 3. Libralces gallicus ? Skull of a young specimen, dredged off the Dogger Bank.

B.M. (G.D.), 46108. | natural size.

GEOL. II, I. 2

i8

DEER OF WEYBOURN CRAG AND FOREST BED OF NORFOLK

to the same species. It was dredged off the Norfolk coast and is highly mineralized,
like the other fossils from the Weybourn Crag and Forest Bed. This brain case
is smaller than the skull from East Runton, and nearly of the same size as the
holotype. The sutures are open, giving evidence that it belonged to a young animal.
The correlation between the antlers, the brain cases and the teeth is made possible
by comparison with the holotype. Two incomplete lower jaws from East Runton

B

FIG. 4. A. Libralces minor, holotype. Lower dentition, no locality. B.M. (G.D.),
M. 6227. B. L. gallicus, lower dentition, East Runton. B.M. (G.D.), M. 6206.
Natural size.

(Figs. 4B, 10 D; Azzaroli, 1952, pi. 15, figs. 3, 4) and some isolated lower molars
from Sidestrand may be attributed to this species.

A fine lower jaw in the private collection of Mr. J. E. Sainty of West Runton,
recorded also above (section " Geology "), was extracted in situ from the Weybourn
Crag between East Runton and West Runton .

The identification of more imperfect specimens from other localities will be dis-
cussed later.

DEER OF WEYBOURN CRAG AND FOREST BED OF NORFOLK 19
MEASUREMENTS : —

E. Runton. Dogger Bank.

B.M. (G.D.) M. 6101 B.M. (G.D.) 46108
Skull :

Occipital breadth . 164 . 145

Occipital height ... 107 . 99

Minimal frontal breadth ca. . 212

Breadth of condyles . 82 . 69

Parietal breadth behind the

pedicles ... 109 . 105

E. Runton
B.M. (G.D.) M. 6206
Lower tooth-row :

Total length .... 136

Length of the three molars . 80

Breadth of M2 . . . . 19

Libralces reynoldsi n. sp.

(Figs. 5 B, 6-8, 9 c.)

1891. Alces sp.f Gunn, pi. 5, fig. 4.

1934. Alces latifrons (Johnson) : Reynolds (pars), fig. 6c.

HOLOTYPE. B.M. (G.D.) M. 6553: a nearly complete brain case, with the left
beam (Figs. 6, 9) figured also by Reynolds, 1933, fig. 6c. Mundesley.

ADDITIONAL SPECIMENS. In the British Museum: two very imperfect antlers,
from Sidestrand and Mundesley; an imperfect humerus from Overstrand; a navi-
culocuboid from Overstrand; two phalanges, from Mundesley and Trimingham;
a fragment of a lower jaw from Overstrand. In the Norwich Museum: a lower
jaw, from Cromer; two imperfect antlers, the one from Trimingham (Gunn, 1891,
pi. 5, fig. 4), the second dredged off the Norfolk coast.

DIAGNOSIS. A much larger species of Libralces than L. gallicus, remarkably
exceeding also the size of Cervalces scotti. Skull and antlers more heavily built
and relatively broader than in L. gallicus', other characters closely similar.

DESCRIPTION. Libralces reynoldsi differs from L. gallicus in its much larger
size, and in some features obviously correlated with this character. The skull is
similarly broad and depressed, and is distinguished by the stronger development
of the supraoccipital and supratemporal crests ; the latter have developed a distinct
knob behind each pedicle. A close comparison between the skulls of L. reynoldsi
and of L. gallicus shows that their proportions are not identical. The skull of the
larger species is relatively broader: it exceeds that of the genotype by roughly
I in length and height, and by ^ in breadth. The holotype of L. reynoldsi seems
to have been an unusually large specimen ; the diameter of its beam above the burr
is 96 mm. In comparison with the strongest specimen of L. gallicus from East
Runton (52 mm.), this gives a cross-section three times as large. Apart from the
strong development of the supraoccipital crest, no external features intended to
counteract the weight of the antlers are seen in the skull, but the thickness of the

20 DEER OF WEYBOURN CRAG AND FOREST BED OF NORFOLK

frontals, which are massive, measures 40 mm., exactly double that in the larger
specimens of L. gallicus. The brain cavity is therefore remarkably small. In the
holotype it is partly filled with the matrix, a hard sandstone impregnated with iron
oxides.

FIG. 5. A. Libralces gallicus, right antler from East Runton. B.M. (G.D.), M. 6554.
B. L. reynoldsi, right antler from Mundesley. B.M. (G.D.), Savin 2223. J natural
size.

The remains of the antlers are very incomplete. Unless too much worn, they
show a very strong burr and a deeply grooved surface; the beam is gently bent.
In the holotype and in the antler of Fig. 5 B, which are the largest specimens I
know and presumably belong to the same individual, the beam is short and the

DEER OF WEYBOURN CRAG AND FOREST BED OF NORFOLK 21

FIG. 6. Libralces reynoldsi holotype, Mundesley. B.M. (G.D.), M. 6553. A, front view ;
B, upper view ; c, lateral view (the pedicle has been broken off and the anterior part of the
brain cavity is exposed). J natural size. (Specimen figured by Reynolds, 1933, fig. 6c).

22 DEER OF WEYBOURN CRAG AND FOREST BED OF NORFOLK

flattening of the palmation is distinctly felt at 25 cm. from the burr. The fragments
from Sidestrand and from the Norfolk coast are nearly of the same size, but very
incomplete. The smaller antler, from Trimingham (Gunn, 1891, pi. 5, fig. 4), has
a diameter of 70 mm. above the burr, and shows that the more slender specimens
tended to develop a long beam, as in L. gallicus: the flattening is not felt until
35-40 cm. from the burr.

The palmation is not preserved among the fossils of East Anglia, but fairly com-
plete antlers, very probably of this species, have been found in Germany (see below).
A maximal span of m. 2 • 50 has been recorded.

The identification of other specimens is made possible by their size.

It may be safely assumed that the muzzle of Libralces displayed the same positive
allometry as the other ruminants. We can expect therefore that the .difference
in size between the teeth of L. gallicus and those of L. reynoldsi is greater than
between their brain cases. No doubt such a comparison is valid only between
broad limits, as it has to allow for individual and sexual variations, but it can be
used here, where the species under consideration differ very greatly in size. Only
two imperfect lower jaws, from Cromer and from Overstrand (Fig. 7), can be
attributed to L. reynoldsi. Their size exceeds that of L. gallicus by roughly \.
The teeth are brachyodont, and display all the characteristic features of Libralces.

A humerus showing the characters of the Alcinae (Fig. 8), evidently belongs to
this species. The proximal epiphysis and half of the distal epiphysis are missing,
but from the remaining part a total length of 445-450 mm. can be inferred. A
naviculocuboid (Fig. 8) and two phalanges may also be attributed to this species.
The naviculocuboid is very large and massive, and relatively higher than the cor-
responding bone of the giant deer. The three cuneiforms are fused with it. The
phalanges are long and slender as in Alces, but much larger.

OTHER LOCALITIES. Several remains from Eastern and Central Europe, attributed
to Alces latifrons by authors, very probably have to be identified with L. reynoldsi.
Pavlow (1906: 7, pi. i, figs, i, 2) described an incomplete antler and a lower jaw
from Tiraspol (Bessarabia). The teeth, although much worn, show the same size
and characters as those of L. reynoldsi. The antler is intermediate in size between
the antlers from Mundesley and that from Trimingham, and its beam is relatively
long; this seems to confirm the inference drawn from the English specimens that
more slender individuals tend to develop longer beams. Another lower jaw with
the same characters has been figured by Soergel (1923, pi. 2, figs. 5, 6). It was found
in the sands at Mauer, together with other teeth and limb bones, among them a
metatarsal of 450 mm. total length. This, too, fits very well with the size of the
humerus from Overstrand. The same author stated (1913, 1923) that an " Alces
latifrons " of nearly the same size is rather common at Mosbach, whereas at
Siissenborn an " Alces latifrons " of a still larger size has been found. But it is
hard to imagine how an animal of this structure could have developed a size still
larger than that of our species. Possibly Soergel's statement was based on com-
parisons between the antlers. According to Vaufrey (1931 : 538) more specimens of
"Alces latifrons " have been found recently at Mosbach. The skull reproduced
in his fig. 5, although too much reduced to permit exact comparisons, seems to belong

DEER OF WEYBOURN CRAG AND FOREST BED OF NORFOLK 23

24 DEER OF WEYBOURN CRAG AND FOREST BED OF NORFOLK

to L. reynoldsi. This author recorded a maximal span of m. 2-50 among the fossils
from Mosbach.

Recently I have seen a fine lower jaw from Mosbach in the Museum of Natural
History at Basel (D. 228), which in all its characters is identical with those of L.
reynoldsi from the Forest Bed.

FIG. 8. A. Libralces reynoldsi, right humerus, Cromer. B.M. (G.D.), M. 6464. J
natural size. B & c. L. reynoldsi, left naviculocuboid, Overstrand. B.M. (G.D.),
M. 6526. ± natural size.

THE RELATIONSHIPS OF Libralces reynoldsi. Although the record of L. reynoldsi
is not fully satisfactory, its affinities with L. gallicus are clearly seen. It is highly
probable that L. reynoldsi is its direct descendant. These two species differ mainly
in size, and increase in size is a common evolutionary trend among ungulates. The
antlers of L. reynoldsi seem to have evolved following a trend already hinted in

DEER OF WEYBOURN CRAG AND FOREST BED OF NORFOLK 25

26 DEER OF WEYBOURN CRAG AND FOREST BED OF NORFOLK

L. gallicus, that is, increase in size together with a shortening of the beam. Other
differences in the skull, as shown above, are evidently correlated with the increase
in weight of the antlers.

L. reynoldsi is distinctly larger than Cervalces scotti, and is the largest cervid so
far known. It may be inferred that it stood m. 1-90 to m. 2-00 at the withers;
its skull was very heavily built, but its limb bones were long and slender as in
the other Alcinae.

MEASUREMENTS : —

Skull of the holotype :

Occipital breadth . . . . . 218

Occipital height . . . . . . 135

Minimal frontal breadth . . . . 278

Breadth of the brain case behind the pedicles . 171

Breadth of the condyles . . . . 132

Lower tooth row (Cromer, Norwich Museum. 2. 116.22)

Length of the three molars . . . . 112
Breadth of M2 . . . . . . 25

Humerus (Overstrand, B.M. (G.D.) M. 6462)

Total length ...... 445-450 (inferred)

First anterior phalanx (Mundesley, B.M. (G.D.) M. 6538)

Total length . . . . . . 100

Proximal breadth ..... 42

First posterior phalanx (Trimingham, B.M. (G.D.) M. 6535)

Total length ...... 101

Proximal breadth ..... 38

Naviculocuboid (Overstrand, B.M. (G.D.) M. 6526)

Antero-posterior diameter .... 75

Transverse diameter ..... 80

Height 42

Libralces cf. reynoldsi

From the Upper Freshwater Bed, West Runton.
(Fig. 7 B.)

A fragment of a lower jaw with the three molars, from the Upper Freshwater
Bed at West Runton, is distinguished by the very poor development of the basal

27

columns. The groove on the anterior crescent of Mx is well marked ; a shallow groove
is present also on the anterior crescents of M2 and M3.

Lib r alee s minor n. sp.

(Figs. 4 A; 10 A, E, F.)

1934. Alces latifrons (Johnson): Reynolds (pars), fig. 6a ?, 7.

HOLOTYPE. A left lower jaw (B.M. (G.D.) M. 6227. No exact locality). Figs.
4 A, 10 A, 10 F.

ADDITIONAL SPECIMENS. Five incomplete lower jaws (one figured by Reynolds,
fig. 7), and a lower molar, from East Runton and Sidestrand.

Possibly some fragments of antlers of a rather small size, from East Runton.
The most complete of these fragments was figured by Reynolds, fig. 6a. Very
doubtful specimens come from West Runton, Overstrand and Trimingham. (See
also below.)

DIAGNOSIS. A Libralces with teeth smaller and distinctly narrower than those of
L. gallicus.

DESCRIPTION. The teeth need no particular description. Apart from the
smaller size and narrower form, they correspond in all respects to those of L. gallicus.
The groove on the anterior crescent of Mx is always clearly seen ; it is not very clear
in Reynold's photograph, which seems to have been retouched. A similar groove
in M 2 is present in three specimens. The cross-section of the mandibular ramus is
also distinctly narrower than in L. gallicus (Fig. 10).

The identification of the antlers is based on their size and is uncertain ; it cannot
be excluded that the specimens tentatively attributed to L. gallicus are young indi-
viduals of other species. The antler figured by Reynolds differs from those of L.
gallicus in its smaller size, its much shorter beam and its relatively larger palmation.
The other specimens from East Runton show the same characters but are still less
complete. One of them (B.M. (G.D.) M. 6550), in Savin's opinion (note in the manu-
script catalogue) is possibly the symmetrical part of Reynolds' specimen.

The fragments of beams from Overstrand, Trimingham and West Runton are
very imperfect and much worn.

MEASUREMENTS : —

Holotype. East Runton.

B.M. (G.D.) M. 6227 B.M. (G.D.) M. 6210
Lower tooth-row :

Total length .... 128 . 125

Length of the three molars . 74 . 71

Breadth of M2 .... 14 . 15

Libralces latifrons (Johnson)
(Figs. 10 B, c ; u)

1874. Cervus latifrons Johnson, pi. i.

1887. Alces latifrons (Johnson) Dawkins (pars), pi. i, fig. 6.

28

DEER OF WEYBOURN CRAG AND FOREST BED OF NORFOLK

HOLOTYPE. A left antler from Happisburgh (Norwich Museum).

ADDITIONAL SPECIMENS. None surely identified, but possibly three lower jaws,
from the Forest Bed of Mundesley (Figs. 10, n), Walcott (Norwich Museum),
and Cromer (private collection of J. E. Sainty). For more doubtful specimens see
also later.

FIG. 10. A. Libralces minor, holotype. B.M. (G.D.), M. 6227. J natural size. (See
also Fig. 4.) B. L. latifrons ? Lower jaw from Mundesley. B.M. (G.D.), Savin 168.
J natural size. C-F. Cross-sections of lower jaws below the hinder lobus of M2.
f natural size, c. L. latifrons? B.M. (G.D.), Savin 168. D. L.gallicus, East Runton.
B.M. (G.D.), M. 6229. E. L. minor, East Runton. B.M. (G.D.), M. 6210. (Speci-
men figured also by Reynolds, 1933, fig. 7.) F. L. minor, holotype. B.M. (G.D.),
M. 6227.

DEER OF WEYBOURN CRAG AND FOREST BED OF NORFOLK 29

DESCRIPTION. This species is based on an incomplete left antler with a part
of the frontal, and is ill denned. The pedicle is long and set horizontally, and the
frontal is very thick, as in the other species of Libralces. The size is intermediate
between those of L. gallicus and of L. reynoldsi. The antler differs from these
species in having a short and straight beam; the palmation seems to have been
very broad. The remaining portion of the frontal gives evidence of a larger animal
than L. gallicus', on the other hand, the shortness of the beam makes it improbable
that this fragment belonged to a young specimen of L. reynoldsi, but this evidence is
by no means conclusive.

In the collections I have seen there are also three lower jaws of a size intermediate
between that of L. gallicus and that of L. reynoldsi. L. latifrons is the only species

FIG. ii. Libralces latifrons? Lower dentition, Mundesley. B.M. (G.D.), Savin 168.

Natural size. (See also Fig. 10.)

with which I can tentatively identify them. The jaw from Mundesley (Figs. 10,
n) is fairly complete. Its cross-section is remarkably narrow and deep. There is
a distinct burr behind the third lobe of M3. A jaw from Walcott (Norwich Museum)
and one found on the beach between Cromer and the Runtons (private collection
of J. E. Sainty) are more imperfect.

MEASUREMENTS : —

Mundesley.
B.M. (G.D.) Savin 168

Lower tooth-row :

Total length

Length of the three molars

Breadth of M,

88

20

30 DEER OF WEYBOURN CRAG AND FOREST BED OF NORFOLK

Lib r alee s incertae sedis

(FigS. 12 A, B, C.)

As stated above, the distinction between the four species of Libralces is based
mainly on the size. Many fragments therefore cannot be identified with certainty ;
in some instances it is practically impossible to distinguish the smaller species from
young individuals of the larger species. A brief account of the most interesting
fragments will be given here.

The fragments figured by Dawkins (1887, pi. i, figs. 2, 3), both much worn, may
possibly belong to L. gallicus. His pi. I, fig. 4 is either L. latifrons or, more probably,
L. reynoldsi. Gunn's type of Alces bovides (1891, pi. i A)may be either L. gallicus or
L. minor. On his plate 5 Gunn figured many specimens attributed to Alces bovides
and Alces sp. The specimen of fig. 5, much rolled, seems to belong to a small species ;
fig. i is the specimen figured by Dawkins on his pi. i, fig. 2; fig. 2 is neither Alces
nor Libralces; fig. 3 is Libralces gallicus (Dawkins' pi. i, fig. 5), and fig. 4 is L.
reynoldsi (see above).

The fragment figured by Reynolds, 1933, fig. 66, is very doubtful, and might
possibly belong to Euctenoceros.

Among the hitherto undescribed specimens, the following are the most inter-
esting :

Fig. 12 A: A fragment of a frontal with the base of the antler, showing a very
strong development of the burr. It belonged to a young specimen, probably L.
gallicus. The frontal is rather thin (14 mm.).

Fig. 12 B : Another fragment of a frontal, with the base of the antler. The pedicle
is very long and the flattening begins near to the burr. This is a young specimen
of either L. latifrons or L. reynoldsi.

B.M. (G.D.) Savin 417. — A skull roof of a young specimen, very probably L.
reynoldsi, from the Forest Bed at Sidestrand. The median suture is open and
forms a median ridge. The supratemporal crests are well developed, the pedicles
small, giving evidence that the antlers were not yet fully developed. In conse-
quence, the frontal is very thin. Its thickness is 27 mm. on the median ridge, but
on the sides of the ridge it is only 16 mm. and the thickness of the parietal is n mm.

Fig. 12 c. — An antler from the Forest Bed at Mundesley evidently belongs to
an abnormal specimen. It bears a small anterior tine on its proximal portion and
shows an abrupt bending at the middle. The characters of the surface sculpture,
which shows very deep and large grooves, makes it probable that this fragment
belongs to Libralces.

Genus CAPREOLUS Hamilton Smith

There is not much to say about this genus. As far as I know, only Capreolus
capreolus has been recorded from the Middle Pleistocene. According to Soergel
(1923) this species is fairly common in Germany. The specimens from Mauer,
which are of a larger size than the living species, have been distinguished by this
author as C. capreolus mut. prisca.

DEER OF WEYBOURN CRAG AND FOREST BED OF NORFOLK 31

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Capreolus capreolus (L.)

1882. Cervus capreolus L. : Newton, p. 52.

1933. Cervus capreolus L. (= Capreolus caprea Gray) : Reynolds, p. 34, fig. 15.

DISTRIBUTION. Forest Bed at Overstrand, Ostend, Sidestrand. Doubtful at
Happisburgh and Cromer. Upper Freshwater Bed at West Runton.

Newton (1882) questioned the occurrence of the roe in the Forest Bed series. In
his opinion a fine antler from Happisburgh (Geol. Survey) may have been found
either in the Forest Bed or in the overlying postglacial drift, whereas another specimen
from Cromer (Baker coll.) is also of doubtful origin. But Reynolds recorded this
species from the Forest Bed of Bacton, Ostend, Overstrand and West Runton (U.
Freshwater Bed ?), and figured three fine antlers. In the Savin collection there are
in addition a lower jaw from the Forest Bed of Sidestrand and several fragments
from the Upper Freshwater Bed of West Runton, namely, two young antlers,
three incomplete lower jaws and several isolated incisors and upper molars. There
is no reason therefore to question its occurrence in the Forest Bed and Upper Fresh-
water Bed. The size is somewhat variable, but the material is too scanty for good
comparisons.

Genus CERVUS L.

The classification of the many species and subspecies of this genus is a thorny
problem to palaeontologists. In the very abundant literature many names have
been proposed, but the value of a great part of them is questionable. The dis-
tinctions have generally been based on antler characters, which may be deceptive
owing to the wide range of individual variation. However, they still give the
most useful data. The remains of the skull are largely represented by the less
significant parts, namely brain cases, lower jaws and teeth. Moreover comparisons
with the living species and races are made difficult by the fact that zoologists base
their classification mostly on external characters, generally giving little attention
to the skulls. However, Pocock's recent studies (1942-430) on the Indian deer
have shown that skulls also may eventually afford valuable data.

A fairly complete account of the species and subspecies of this genus has been
given by Lydekker (1915: 116 ff.). He distinguished in the genus Cervus seven
species; on one of these, C. albirostris, Flerow (1930) has founded the genus
Przewalskium. The remaining six species are: C. elaphus L., C. canadensis Erxl.,
C. yarkandensis Blanford, C. wallichi Cuvier, C. macneilli Lydekker, C. cashmirianus
Gray; the last falls under the synonymy of C. hanglu Wagner. They are all char-
acterized by the presence of a bez tine.

C. canadensis, the wapiti, with many varieties ranging from N. America to Central
Asia, and C. macneilli, ranging over Szechuan, Kansu and Yunnan, are distinguished
by the upper tines being placed in a plane parallel to the median plane of the head.

In the other species the upper bifurcations take place transversally to the median
plane. C. wallichi and its variety C. wallichi affinis, the shou (Sikkim and Tibet),
are of a very large size. The antlers have generally five tines and end with a simple

DEER OF WEYBOURN CRAG AND FOREST BED OF NORFOLK 33

transversal fork. An accessory tine seems to occur very rarely. This species has
been redescribed recently by Pocock (1942-43^), who gave the two varieties full
specific rank. Cervus hanglu, the hangul, from Kashmir, is a little smaller than
C. wallichi. Its antlers are similarly built, but an accessory tine or even a double
terminal fork are not uncommon (see the good photograph by Stockley, 1948).
This species also has been redescribed by Pocock (1942-43^). C. yarkandensis,
from E. Turkestan, has antlers of the same type, generally with five tines, eventually
with an accessory tine.

C. elaphus, the familiar red deer, is distinguished by a greater complication of the
antlers. As a rule they end with four or more tines forming the characteristic
" cup " (" Krone " of the German authors), originated by the fusion of two or
more transversal forks (Beninde, 1940). The cup is very variable, especially in
some living stocks, which in many ways are influenced by unnatural conditions of
life. This, however, does not alter the fundamental fact that the cup is a peculiar
feature of Cervus elaphus. In the Upper Pleistocene European red deer it is fairly
constant.

This species has given origin to many varieties or geographical races ; the eastern
races are generally larger, but do not attain the size of the wapiti. In the smaller
races the antlers are less complicated and the bez tine is sometimes lacking, but the
cup, as a rule, is still present.

The typical area of the species is Southern Sweden. Lydekker (1915) listed the
following subspecies: C. elaphus barbarus Bennett, Algeria and Tunisia; C. elaphus
corsicanus Erxl., Corsica and Sardinia; C. elaphus hispanicus Hilzheimer, S.W.
Spain; C. elaphus atlanticus Lonnberg, Trondhjem district, Norway; C. elaphus
scoticus Lonnberg, Scotland; C, elaphus hippelaphus Kerr, Germany; C. elaphus,
unnamed subspecies, Bukovina; C. elaphus maral Ogilby, Caucasus and Persia.
To these we may add C. elaphus bolivari Cabrera, Central and Northern Spain;
C. elaphus brauneri Charlemagne, Crimea.

The value of some of these subspecies, even as geographical races, is questionable.
The craniological studies by Ingebrigtsen (1922-23, 1927) are instructive. According
to this author, no distinction is possible between the red deer of Sweden, Germany
and the Norwegian mainland districts. Some stocks of the Norwegian isles, especi-
ally from Hitra, have developed a smaller size, which is due only to environmental
factors and is no expression of hereditary differences. This is true also for the
Scottish red deer (see also Ritchie, 1920), and is possibly true for C. elaphus corsicanus
and C. elaphus hispanicus. But two subspecies at least seem to be genetically estab-
lished : the small C. elaphus barbarus, distinguished by its spotted pelage, and the
large C. elaphus maral, with large but relatively simple antlers and cruciform nasals;
this character, as far as I know, has not been recorded in the literature, but is well
developed in all the skulls of Caucasian stags exhibited in the British Museum.

No attempt at a complete revision of the fossil red deer will be made here, but it
may be useful to give a short account of its earlier representatives.

The red deer has no relatives in the Villafranchian faunas of Central and Western
Europe. Redstone (1930) quoted it from the Red Crag, but Dr. K. P. Oakley
informs me that the specimen referred to actually came from overlying late Pleisto-

GEOL. II, I. 3

34

cene deposits. The red deer immigrated from Eastern Europe or from Asia during
the Mindel-Riss interglacial, and possibly other immigrations took place later. The
best known representatives of the oldest European red deer are not identical with
living red deer, and there is no proof of the occurrence of the typical form of Cervus
elaphus in the Mindel-Riss interglacial.

The most perfect representatives of the genus Cervus s. str. were found at Mosbach
and Mauer and have been described by Beninde (1937). Further details were
added by Haupt (1938), Kleinschmidt (1938) and Schmidtgen (1938).

Cervus acoronatus Beninde, from the sands of Mosbach (main fauna), is a species
of large size. It has stout and massive antlers, with five tines, namely, well developed
brow and bez tines, a trez tine and a flattened terminal fork, set transversally to
the axis of the body. Schmidtgen (1938) has figured an unusually flattened terminal
fork, and Kleinschmidt (1938) a terminal fork with a small accessory tine. As
a whole, C. acoronatus seems to constitute a rather uniform and well characterized
stock. The lack of the cup led Beninde to distinguish it from Cervus elaphus as
a full species. The antlers strikingly recall those of the red deer of Central Asia,
especially of the large C. wallichi, from which they seem to differ only by the marked
flattening of the upper fork and by a stouter form.

Haupt (1938 : 32) proposed to change the name C. acoronatus into C. elaphus
mosbachensis, but this is obviously contrary to the rules of nomenclature. Ac-
cording to Beninde, imperfect antlers of a smaller size occur in the same sands. It
is not clear whether they belong to young individuals or to a distinct species.

The deer from Mauer, of nearly the same age if not a little younger, is more poorly
represented; it has no typical cup and closely resembles C. acoronatus, from which
it is distinguished by a twisting of the upper fork, bringing its anterior surface
obliquely outwards, and by the presence on it of accessory tines, eventually lying
out of its main plane.

Soergel (1923) gave this species the name C. elaphus mut. prisca, changed into C.
elaphus priscus by Beninde, but this name is preoccupied. Kleinschmidt (19380)
proposed the name C. benindei, and in the same year Haupt proposed C. elaphus
mauerensis. C. benindei has the priority, as it was published in the first months
of 1938 (the fascicle was purchased by the British Museum in March), whereas
Haupt 's paper was published in the second half of the year (see on p. 3 of the same
volume the necrology of G. Klemm, who died on 6th August, 1938).

This form is more closely related to C. acoronatus than to later red deer; there is
no reason therefore to rank it as a subspecies of C. elaphus Here it will be called
C. benindei.

In 1937 Beninde supposed that the upper fork of the red deer from Mauer might
represent an intermediate stage, leading to the formation of the cup. This however
is not in accordance with his later conclusions (1940) on the origin of the cup, based
on comparative studies of recent red deer, that is, that the cup originated through
the fusion of two or more transversal forks. The peculiar position of the terminal
fork of C. benindei does not occur in later red deer. Beninde himself seems to
have changed his mind, as he wrote (1940 : 256) : " Im I Interglazial des Oberrheins
(Mosbach, Mauer) liegen Geweihe, die gegen den heutigen Rothirsch eine ganz

DEER OF WEYBOURN CRAG AND FOREST BED OF NORFOLK 35

eigene Stellung einnehmen. Aber bereits um das Riss (Steinheim a.d. Murr, Primi-
genius-horizont) erscheinen Hirsche vom rezenten Typus . . .

The validity of C. acoronatus and C. benindei has been questioned by Bachofen
Echt (1941), who claimed to see intermediate stages between these two species and
C. elaphus among the fossils from Hundsheim. However, his observations, which
are based on two imperfect specimens, do not invalidate Beninde's conclusions.
Moreover the age of Hundsheim, according to Kormos (1937) is " ... vielleicht
etwas j linger als das Upper Freshwater Bed von West Runton." If so, it is younger
than the main faunas of Mosbach and Mauer.

The remains from the other localities of the same age are scanty and imperfect.
They are not identical with the species of Mosbach and Mauer, but do not demon-
strate the presence of the true Cervus elaphus in the second Interglacial. The
denomination Cervus cf . elaphus better expresses our state of knowledge.

The red deer from Siissenborn, to which Pohlig (1909) gave the name C. elaphus
trogontherii, is rather large, and seems to be characterized by the lack of the bez
tine. The status of the red deer in this locality is however not clear. Some years
later Soergel (1923 : 221) quoted Cervus elaphus cf. mut. prisca Soerg., Cervus maral
fossilis Ogilby (sic), Cervus (Elaphus n. sp., grosse Form).

A basal portion of an antler from Tiraspol (Pavlow, 1906, pi. i, fig. 8) is large and
stout and bears a strong bez tine.

The red deer from the Upper Valdarno (Azzaroli, 1948) also belongs to this period.
It is of a medium size and its antlers are poorly known.

Stehlin (1932) made reference to other early red deer from Jockgrim, Bammental,
St. Prest and Solilhac. The faunas of Solilhac however are not older than the
Riss glaciation (Bout & Cailleux, 1951).

The true Cervus elaphus is certainly present in the Riss glaciation. Good speci-
mens from the Antiquus-horizon of Steinheim a.d. Murr have been distinguished
by Beninde (1937) under the subspecies C. elaphus angulatus. The red deer of the
overlying Primigenius-horizon is not distinguishable from the typical form.

Cervus cf. elaphus L.

(Figs. 13, 14.)

1882. Cervus elaphus ? L. : Newton, p. 55.

1891. Cervus elaphus L. : Newton, p. 27, pi. 4, fig. 14.

1931. Cervus elaphus L. : Reynolds, p. 4.

OCCURRENCE. This is a very common species in the Forest Bed and in the Upper
Freshwater Bed, but its remains are rather poor. In 1882 Newton questioned the
occurrence of the red deer in the Forest Bed, but included it in his faunal list in
1891. He pointed out that it was then the earliest record of the red deer. Reynolds
recorded it from the Forest Bed at Happisburgh, Overstrand, East Runton and
Trimingham. The specimens I have seen bear the indications : Trimingham,
Sidestrand, Bacton, Overstrand, Happisburgh, Mundesley, East Runton (one
specimen, much rolled), West Runton (Upper Freshwater Bed), Palling (on the
beach).

36 DEER OF WEYBOURN CRAG AND FOREST BED OF NORFOLK

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DESCRIPTION. The antlers consist for the most part of basal fragments, and
differ greatly in size; some of them are very massive (Fig. 13). The bez tine is
generally well developed, and in some instances is pushed upwards. In one young
specimen it does not occur in its typical form, but we find here a small tine which
may be interpreted indifferently as a bez or a trez (Fig. 13 D). Possibly however
some fragments of this species without a bez have escaped my attention.

The only fairly complete antler is of medium size and is very stout. Its upper
portion ends with three tines originating from two successive bifurcations, and is
damaged. This antler cannot be identified with the species of Mauer and Mosbach,
but does not show the typical cup of C. elaphus. As it seems to belong to a not
fully grown specimen the question of its identity is still open.

FIG. 14. Cervus cf. elaphus, lower dentition, Trimingham. B.M. (G.D.), M. 6220. Natural

size.

Four lower jaws, from Trimingham and from the Upper Freshwater Bed at
W. Runton, do not show any remarkable feature. P4 is always advanced.
The attribution of some limb bones will be discussed later.

Genus EUCTENOCEROS Trouessart

A definition of this genus and a discussion of its synonymy have been given in
previous papers (Azzaroli, 1948; Bout & Azzaroli, 1953). In these papers I did
not include in it " Cervus " falconeri Dawk., whose relationships were not clear to
me. Kunst (1937) pointed out its affinities to E. teguliensis (= E. ctenoides; see
below), but as a matter of fact these two species differ in many characters. On
the contrary its affinities with E. sedgwicki are very close. I therefore include it
in this genus. It is the oldest and most primitive of its species, but not their
common ancestor.

Unfortunately in 1948 I overlooked Nesti's description of the species from the
Upper Valdarno, which is not quoted in Forsyth Major's papers and is recorded
only in Sherborn's catalogue. Nesti exhibited the deer from the Upper Valdarno
at a congress of Italian scientists held in Florence in 1841, and proposed for the
larger species the names Cervus dicranios, Cervus ctenoides and Cervus orticeros.
His verbal description was summarized in the report of the meeting by the secretaries,
Savi & Sismonda (1841). The name Cervus orticeros has been subsequently ignored
and is probably synonymous with C. ctenoides.

These two species have been briefly described also by Cornalia (1858-71: 61).
The incorrect spelling dicranius has been introduced by Forsyth Major.

The species of this genus display very peculiar features in the antlers, which make

38 DEER OF WEYBOURN CRAG AND FOREST BED OF NORFOLK

them easily distinguishable from other genera. The first tine branches from the
outer anterior part of the beam ; the following tines arise from its anterior part and
lie in a plane. They are numerous, three in E. falconeri, four or five in the other
species, and are frequently more or less undulated. Beam and tines display a
tendency to become flattened and in some species the tines subdivide further. The
flattening is generally more pronounced in the upper portion of the antlers, but in
E. sedgwicki reaches as far down as the brow tine. In the larger specimens of some
species a small accessory tine, directed upwards, is given off from the first bifurca-
tion.

With the sole exception of E. falconeri all the species of this genus are of a large
size. Together with the common features described above, they display a remark-
ably wide range of variation which makes specific distinctions difficult. I do not
feel satisfied with some of the determinations I published in 1948. An antler from
Olivola (fig. 8) then attributed to E. ctenoides is actually of E. dicranios. The
tines of this specimen do not show the strong backward bending of the lectotype,
but in its general characters this antler is more similar to it than to E. ctenoides.
Moreover, my reconstruction was wrong: the lower tine preserved in this antler
corresponds to the second tine, not to the third tine of the lectotype of E. dicranios.

E. ctenoides and E. teguliensis are very probably identical. The antlers do not
afford any good basis for distinction.

Eiictenoceros tetraceros (Dawkins)
(Figs. 15, 16, 17 A, B, D.)

1878. Cervus tetraceros Dawkins, p. 416, figs. 14, 17.

1891. " Cervus tetraceros Mackie " : Newton, p. 32, pi. 4, fig. 2.

1953. Euctenocevos tetraceros (Dawk.) : Azzaroli in Bout & Azzaroli, p. 43, figs. 1-6.

OCCURRENCE. Fairly common at East Runton, rare at Overstrand.

DESCRIPTION. This species is represented by two fine antlers (Fig. 15) and several
fragments. They all agree with the specimens from Peyrolles; one specimen
only (Fig. 16) is distinguished by the abnormal inward bending of its brow tine.
As at Peyrolles, all the antlers of this species have been naturally shed.

Three lower jaws show the same characters as those from Peyrolles. Probably
also a fourth lower jaw with a primitive P4 belongs to this species.

MEASUREMENTS : —

East Runton. East Runton. East Runton.

Lower tooth rows : B.M. (G.D.) M. 6232 B.M. (G.D.) M. 6218 B.M. (G.D.) M. 6221

Total length . . . 131 . 134 . 133

Breadth of M2 . 16 . 15 . 15

Euctenoceros ctenoides (Nesti)

1841. Cervus ctenoides Nesti (in Savi & Sismonda, p. 159).
1858-71. Cervus ctenoides Nesti : Cornalia, p. 61.

DEER OF WEYBOURN CRAG AND FOREST BED OF NORFOLK 39

B

FIG. 15. Euctenoceros tetraceros, East Runton. J natural size. A. Left antler, B.M.
(G.D.), M. 6369. B. Right antler, B.M. (G.D.), M. 6370.

40 DEER OF WEYBOURN CRAG AND FOREST BED OF NORFOLK

1887. Cervits savini Dawkins (pars), pi. 3, fig. 4.

1948. Cervus (Euctenoceros) ctenoides Nesti : Azzaroli (pars), p. 62, figs. 6, 7, 9.

1947. Eucladoceros teguliensis Dub. : Hooijer, p. 34 (cum syn.).

SYNONYMY. The distinction between E. ctenoides and E. teguliensis does not
seem to be valid. The ranges of variation of these two species widely overlap.
The antlers from Tegelen do not attain the size of the larger specimens from the
Upper Valdarno, but this may be due merely to the incompleteness of the record.
The dentition and limb bones are of the same size.

FIG. 1 6. Euctenoceros tetraceros, abnormal right antler, East Runton.
B.M. (G.D.), M. 6399. J natural size.

Among the specimens from East Anglia a very fine antler from Sidestrand
(Dawkins' pi. 3, fig. 4) so closely resembles the lectotype of E. ctenoides that no one
would hesitate to identify it with this species.

OCCURRENCE. The only good evidence of the occurrence of this species is given
by the antler from Sidestrand figured by Dawkins, who incorrectly attributed it
to Cervus savini, and by some basal fragments from East Runton, of a slightly
smaller size.

For the identification of other fragments of antlers and of dentitions see later.

Euctenoceros sedgwicki (Falconer)

(FigS. l8, IQ, 20 A, 21 A, F.)

1868. Cervus sedgwicki Falconer (ex Gunn MSS.), p. 472, pi. 37, figs. 1-3.

1882. Cervus sedgwicki Falc. : Newton, p. 60.

1891. Cervus sedgwicki Falc. : Gunn, pi. 3, figs. 99, 100.

1891. Cervus sp. Gunn, pi. 3, figs. 105, 106.

1891. Cervus sedgwicki Falconer : Newton, p. 31.

DEER OF WEYBOURN CRAG AND FOREST BED OF NORFOLK 41

OCCURRENCE. This species is fairly common, but is represented for the most
part by very imperfect fragments. The only satisfactory specimen is the holo-
type (Fig. 18). This and most of the other fragments were found at Bacton, a
few specimens at Mundesley and a much rolled fragment at East Runt on. For
doubtful fragments see also below.

FIG. 17. Euctenoceros ; dentition, East Runton. Natural size. A. E. tetraceros,
B.M. (G.D.), M. 6218 (advanced form). B. E. tetraceros, B.M. (G.D.), M. 6221 (primi-
tive form), c. E. ctenoides ? B.M. (G.D.), M. 6241 (primitive form). D. E. tetraceros,
B.M. (G.D.), M. 6206 (advanced form). E. E. ctenoides? B.M. (G.D.), M. 6213.

DESCRIPTION. This species is of a large size and is characterized by the high
position of the brow tine, 10-15 cm- above the burr, and by a very marked flattening
of the upper portion of the beam and of the tines, which divide into three or four
secondary tines. The brow tine is also much flattened, and in the type it divides
into four points. The high position and the flattening of the brow tine are remark-
ably constant, and are well displayed even in young specimens (Fig. 21 F). The
pedicles are set near one another, as in E. boulei and E. dicranios.

42 DEER OF WEYBOURN CRAG AND FOREST BED OF NORFOLK

FIG. 1 8. — Euctenoceros sedgwicki, holotype, Norwich Museum, Gunn Colin.
No. 99. T^ natural size.

FIG. 19. Euctenoceros sedgwicki, reconstruction of the holotype. Norwich Museum,
Gunn Colin. No. 99. J natural size.

DEER OF WEYBOURN CRAG AND FOREST BED OF NORFOLK 43

The upper portion of the type antler is badly damaged and its reconstruction is
rather puzzling. On Fig. 18 I have indicated the most probable position of the
fragments. The small cross-section of the beam at the broken surface makes it
unlikely that there were additional tines. On Fig. 19 I have attempted a recon-
struction of this specimen.

COMPARISONS. Euctenoceros falconeri (Dawkins, 1868; see also Kunst, 1937:
101) is a species of medium size. It does not occur in the Weybourn Crag nor
in the Forest Bed, but has been found in the Red Crag and in the Norwich Crag.

FIG. 20. Euctenoceros sedgwicki and E. falconeri, lower portions of the antlers. £
natural size. A. E. sedgwicki, Mundesley. Norwich Museum 323. B. E. falconeri,
Thorpe, Norwich Crag. Norwich Museum 313. c. E. falconeri, Horstead, Norwich
Crag. Norwich Museum 310.

It occurs also in the Poederlian of the Netherlands and in deposits of the same age
in Belgium. The most complete specimens were found in the Belgian Kempen.
The remains from East Anglia are rather fragmentary ; the most complete are the
holotype (a young individual), and a crushed full-grown antler from the Red Crag,
now in the Museum at Ipswich. In the British Museum and at Norwich I have
seen other fragments from the Norwich Crag, and at Ipswich fragments from the
Red Crag. The remains of the Norwich Crag may possibly have been derived from
the Red Crag. Those from the Red Crag bear no exact indication of the horizon,
but their state of fossilization shows that they were not derived.

Two specimens in the Norwich Museum were figured by Gunn (1891, pi. 2, fig.

44 DEER OF WEYBOURN CRAG AND FOREST BED OF NORFOLK

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DEER OF WEYBOURN CRAG AND FOREST BED OF NORFOLK 45

97 ; pi. 7, fig. 3), who did not determine them. Two other specimens of the same
species were attributed to Cervus carnutorum Laugel and to Cervus ardeus Croizet &
Jobert by Newton (1891 : 25-26, pi. 4, figs. 3, 6).

The relationships between E. falconeri and E. sedgwicki are very close. They
are both distinguished by the high position of the first bifurcation and by the well-
marked flattening of the upper portion of the antlers. Very probably E. sedgwicki
is the descendant of its older ally, from which it is distinguished only by more
advanced characters, namely, the larger size, the greater number of the tines and
their more pronounced flattening.

The relationships with E. boulei (Teilhard & Piveteau, 1930) are also very close.
In E. boulei the flattening is confined to the upper tines; its range of variation
seems to be very wide.

The relationships of E. sedgwicki with E. dicranios from the Upper Valdarno
(Azzaroli, 1948) seem to be less close. The range of variation of this species is little
known ; but it is always distinguished by the low position of the brow tine, by a
peculiar backward bending of the upper tines and by their low grade of flattening.
The tines of its lectotype are more numerous than in the type of E. sedgwicki.

The few specimens from Olivola (see also above, introduction to the genus Euc-
tenoceros) do not agree perfectly with those of the Upper Valdarno. In the more
complete skull (Azzaroli, 1948, fig. 10) the first bifurcation is set 8 cm. above the
burr, and in the antler (fig. 8, incorrectly determined E. ctenoides) the backward
bending of the upper tines is less pronounced. These specimens are probably
slightly older than those from the Upper Valdarno (Azzaroli, 1950). E, dicranios
may also be a descendant of E. falconeri; the question however is still open.
The frontal from St. Prest, figured by Gervais (1867-69, pi. 16, fig. 4) and
determined as Cervus (Megaceros) carnutorum Laugel, might possibly belong to
E. sedgwicki. The name Cervus carnutorum was based on fragments of different
species and should be dropped (Stehlin, 1912).

Euctenoceros, incertae sedis

(FigS. 17 C, E, 21 B-E, G, 22)

Three lower jaws and a maxilla from East Runt on and Sidestrand might be
attributed indifferently to E. ctenoides or E. sedgwicki. E. sedgwicki is not otherwise
recorded with certainty from these localities. The teeth are slightly larger, broader
and more brachydont than those of E. tetraceros; the upper molars bear a dis-
continuous cingulum.

A large basal fragment of an antler from East Runton is rather puzzling (Fig.
21 c). It strikingly recalls E. dicranios from the Upper Valdarno.

Two bifurcated tines from the same locality (Fig. 21 D, E) might perhaps be
attributed to E. ctenoides but are very imperfect

An upper portion of an antler (Fig. 21 G), and two similar fragments, from East
Runton and Overstrand, may be attributed indifferently to E. ctenoides or to E.
sedgwicki (cf. the two antlers of E. ctenoides in the Museum at Basel, figured by
Kunst, 1937 : 42).

46 DEER OF WEYBOURN CRAG AND FOREST BED OF NORFOLK

An antler from Overstrand (Fig. 21 B), with an abnormal outer tine near the base
of the second tine, might also belong to E. ctenoides.

A fragment of a frontal with an abnormally reduced antler from East Runton
(Fig. 22) affords an example of degeneration, probably due to very old age. The
antler is much thinner than the pedicle and is reduced to a forked stump. Among
the species from this locality, E. tetraceros, is, for its size, the only one to which this
fragment might be attributed. Also a skull-roof from the same locality, with shed
antlers (B.M. (G.D.), M. 6300), might belong to E. tetraceros. This specimen is
of little interest. Its supraoccipital crest is weak!

A fragment of an antler attributed to " Alces latifrons " by Reynolds (1933, fig.
6b) might possibly belong to E. sedgwicki.

FIG. 22. Euctenoceros ? sp. Basal portion of a left antler. Abnormal specimen, presumably
very old. East Runton, B.M. (G.D.), Savin 1776. £ natural size.

MEASUREMENTS : —

Lower jaw :

Total length of the tooth row
Breadth of M2

Skull roof —

Occipital breadth
Occipital height .
Minimal frontal breadth ca.
Breadth of condyles

Sidestrand.

B.M. (G.D.) Savin 1615

141

18

East Runton.
B.M. (G.D.) M. 6300

135

84
146

81

Genus MEGACEROS Owen

Many species more or less closely related to the familiar giant deer have been
described in the last decades. They have generally been united under the name

DEER OF WEYBOURN CRAG AND FOREST BED OF NORFOLK 47

Megaceros, alternately taken as a full genus or ranked as a subgenus of Cervus.
Some terminal forms from North Africa and Eastern Asia have been separated
under the subgenera Megaceroides Lydekker and Sinomegaceros Dietrich.

It is easily seen that this group is characterized by peculiar features of the skull,
the antlers, the dentition and the limb bones ; and although it is a plesiometacarpal,
it widely differs from the other Cervinae. It is consistent therefore to give it full
generic rank.

Megaceros is characterized by its large size, heavy build, large antlers, which gener-
ally are more or less palmated, and by a strange hyperostosis of the mandibular
ramus and in a lesser degree of the skull, which eventually leads to monstrous
forms. The significance of this hyperostosis has puzzled many palaeontologists,
who often considered it a mark of degeneration or even a pathological character.
However its occurrence in all the species of this genus gives evidence that it is not
pathological in the common signification of the word; the more so as in the most
widespread species it does not display any particularly wide range of variation.
On the other hand it can hardly be considered a useful character, but always has
a distinct mark of abnormality. Much light has been thrown on this point by
Young's studies on Megaceros pachyosteus (1932), a very abundant species from
the Sinanthropus site at Chouk'outien and other localities of Central China. In
this species, which obviously belongs to a terminal branch, the hyperostosis is
extreme. At the same time it displays a very wide range of variation, both in
degree and in the way it affects the bones, and is accompanied by characteristic
marks of degeneration, namely a shortening of the muzzle, small body size with
an unusually wide range of variation, and the frequent occurrence of distinctly
dwarf individuals. Young came to the conclusion that ' ... in the case of
Cervus pachyosteus we witness a zoological group in some state of high zoological
instability (or ' aff clement '), this condition being due to a factor (the hyperostosis)
which can be called a morbid one, since it was not directed to the formation of useful
features."

Zdansky (1925-27) attributed the hyperostotic bones to various genera, but
according to Young these identifications are incorrect.

Similarly degenerate characters (an extreme degree of hyperostosis, a very marked
shortening of the muzzle and a small size) are displayed by another terminal form,
M. algericus Lydekker (in Arambourg, 1938). This hyperostosis, as will be shown
later, is well developed even in the earliest representatives of this genus in Europe,
and there is no evidence of an increase of it with time.

An attempt to trace the phylogeny of the giant deer was made by Soergel (1927),
who ranged the species in a unique phyletic line and assumed that they were derived
from a Pliocene or Lower Pleistocene ancestor from N. Italy, Cervus pliotarandoides
Alessandri. Soergel based his observations mainly on the antlers, and came to the
conclusion that their evolution, starting from an hypothetical Pliocene ancestor
with two basal tines (which he homologized with the brow and the bez tine of the
red deer) was characterized by the disappearance of the lower (brow) tine and by a
gradual downwards shift of the upper (bez) tine till close to the burr.

Many data have been made known since, and Soergel's conclusions need revision.

48 DEER OF WEYBOURN CRAG AND FOREST BED OF NORFOLK

First of all, a strict homologizing with the brow and bez tine of the red deer is not
possible. The antlers of Megaceros are built on a peculiar plan, very different from
that of Cervus and the other Cervinae. We shall follow here the current nomen-
clature and refer to the fairly constant basal tine as the brow tine. Some species
occasionally develop an inconstant additional tine below it, which will be referred
to as the accessory basal tine. Moreover, the chronological sequence is not in
accordance with Soergel's views.

The species of giant deer recorded in the literature are very numerous. Many
of them, of course, are poorly known and of questionable value, but the better known
species can easily be divided into two groups.

The skulls of the older species consist only of more or less complete brain cases.
The antlers as a rule vary greatly ; both skulls and antlers however give very valu-
able information on the relationships of the species. The teeth afford good examples
of evolutionary progress with time, but seem to have advanced along parallel lines
and are therefore of a more restricted use in classification. Moreover, their identi-
fication with skulls and antlers is sometimes uncertain.

The limb bones are robust. Young pointed out the broad form of the distal
epiphysis of the metapodials in Megaceros pachyosteus. This character, although
less pronounced than in the reindeer, is quite distinct and seems to occur in all the
species of this genus. It has not yet been possible to establish whether limb bones
may give some indication for classification.

The two groups into which this genus can be divided will be called, from their
best known representatives, the group of Megaceros giganteus (= M. euryceros
Aldrov. ; for the priority of the name see Berckhemer, 1941) and the group of
M. verticornis Dawk. The subgenera Megaceroides and Sinomegaceros mentioned
above probably represent terminal branches of each group, but their relationships
with the European species are not known with sufficient detail. Moreover the value
of Sinomegaceros has been questioned by Teilhard (1936). These two subgeneric
names therefore will not be used here.

The Group of Megaceros giganteus

Pedicles divergent, but generally set near one another. Forehead hollowed in
front of the pedicles (Fig. 23 D), and eventually swollen into a prominent transverse
ridge between them. Brow tine branching off closely near the burr, flattened and
more or less expanded, fluctuating in some of the younger species.

To this group I attribute: Megaceros giganteus Blumenbach; M. antecedent
Berckhemer; M. savini Dawkins; M. sp. from Felixstow, Red Crag, East Anglia;
M. pachyosteus Young; M. ordosianus Young; M. flabellatus Young; M. yabei
Shikama.

Megaceros giganteus is the largest and most advanced species of this group, and
is very familiar to palaeontologists. It was very widespread through Europe during
the last interglacial and glacial phases. According to Mitchell & Parkes (1949),
the Irish race was an inhabitant of grassy lowlands and the period of its maximal
spread coincides with a mild Wiirm Interstadial, corresponding to the Allerod

DEER OF WEYBOURN CRAG AND FOREST BED OF NORFOLK

49

stage of Denmark; it seems to have become extinct with the last cold phase. The
races of Central and Southern Europe had presumably the same habits of life. I
have not been able to find satisfactory information on their chronological range, but
there is no evidence that any of them outlived the last glaciation (see also Frentzen
& Speyer, 1928).

FIG. 23. Frontals of Megaceros, about J natural size. A. M. verticornis, Pakefield.
Norwich Museum 365. B. M. verticornis, Kessingland. Norwich Museum 364.
c. M. dawkinsi, Sidestrand, B.M. (G.D.), M. 6302. (See also Fig. 33A.) D. M.
savini, Kessingland. Norwich Museum 321.

GEOL. II, I. A

50 DEER OF WEYBOURN CRAG AND FOREST BED OF NORFOLK

Further references to this species may be found in Cornalia (1858-71), Pohlig
(1892), Pavlow (1906), Hescheler (1909), Fabiani (1919), Frentzen & Speyer (1928),
Reynolds (1929), Zakrewska (1932, 1935), Kunst (1937), Kirchner (1937), Azzaroli
(1948). Pohlig distinguished three races, hiberniae, germaniae and italiae, based on
antler characters. This distinction has not been accepted by all authors. Recently
Kirchner rejected it, claiming that their ranges of variation widely overlap, but I
am unable to accept his views. I grant that the variation is great and that extreme
variants may eventually overlap, but on the whole Pohlig's specimens from Germany
are quite distinct from the Irish specimens, and the distinctive characters listed by
this author hold good, at least in a statistical sense. This is true also for the
specimens described by Frentzen & Speyer. In discussing the range of variation
of the German race Kirchner made reference to an incomplete antler from Bergrhein-
feld a. Main, which actually seems to fit better with the Irish race; but its age is
somewhat uncertain, possibly younger than Pohlig's specimens, and it cannot be
taken therefore as a good representative of the German race.

The distinction of the Italian race is less clear, owing perhaps also to the incom-
pleteness of the record. The identifications with Pohlig's races, made by some
authors for specimens from other districts, are questionable. No doubt more
detailed information on the age of the specimens will throw much light on their
relationships and on the value of these smaller systematic units.

Megaceros antecedent from the antiquus-honzon of Steinheim a.d. Murr (a mild
Riss interstadial ; Berckhemer, 1941) has been described as a subspecies of M.
giganteus, but I prefer to give it full specific rank. It is smaller than M. giganteus
and its dentition is more primitive. The antlers are stout, little divergent and
broadly palmated. The brow tine is expanded into a small palmation. A related
form, not yet well defined, occurs in the overlying primigemus-horizon.

Megaceros savini from the Forest Bed is still more primitive. The brow tine is
widely expanded, the beam is long and flattened and ends with three tines. This is
the only example of the lack of a palmation in the genus Megaceros. The dentition
is primitive. A more detailed description will be given later.

Megaceros sp. ind. from the Red Crag at Felixstow is represented by a basal
fragment of an antler, described and figured by Owen (1856, fig. 18), and is now in
the Museum at Ipswich. It is a very poor fragment. The brow tine is broken;
its base lies close to the burr and is slightly expanded, as in M. giganteus; the
beam shows the same sigmoidal swing. For these reasons I attribute it to this
group. There is no certain information about the horizon of this specimen. Its
fossilization shows that it was derived possibly from the lower part of the Red Crag.

Megaceros pachyosteus (Young, 1932) was found at the Sinanthropus site at
Chouk'outien and at other localities of Central China. At Chouk'outien it is
extremely abundant. Its skull and antlers clearly show the characters of this group ;
it is distinguished by a relatively small size, a very brachyodont dentition, an
extreme hyperostosis of the skull, and by other characters clearly pointing to
degeneration. The skull is very broad and depressed, the muzzle short, the body
size is very variable and distinctly dwarf individuals are not uncommon. Also the

DEER OF WEYBOURN CRAG AND FOREST BED OF NORFOLK 51

antlers are very variable, but are always very stout and broadly palmated. The
brow tine is also palmated. The dentition is still rather primitive.

Megaceros ordosianus from the loess of Manchuria, M. flabellatus from locality 9 at
Chouk'outien and M. yabei from the Upper Pleistocene of Japan (Young, 1932;
Teilhard, 1936; Shikama, 1939) are distinguished from M. pachyosteus by a less
developed hyperostosis and by some characters of the antlers. According to
Teilhard the wide range of variation makes specific distinctions somewhat difficult.

The Group of Megaceros verticornis

Pedicles divergent, generally set wide apart and directed obliquely backwards.
Forehead transversely flat or convex in front of the pedicles (Fig. 23), generally
marked by two faint ridges running along the median bases of the pedicles and
vanishing on the forehead. No transverse ridge on the skull roof between the
pedicles. Brow tine typically subcylindrical, arising from the upper portion of the
beam at some distance from the burr and strongly bent forwards. Very often a
more or less distinct knob below it on the anterior part of the beam, which may
eventually develop into an accessory tine, close to the burr. Some species display
various degrees of reduction of the brow tine.

To this group I attribute Megaceros verticornis Dawkins ; M. aff . verticornis from
Sussenborn; M. aff. verticornis from Trimley, Red Crag; M. dawkinsi Newton;
M. belgrandi Lartet from Montreuil (= M. dawkinsi?}; M. aff. belgrandi from
Taubach; M. aff. belgrandi from Laufen; M. aff. belgrandi from Tiraspol; M.
mosbachensis Soergel ; M. solilhacus Robert ; M. algericus Lydekker ; M . pliotaran-
doides Alessandri ; M. cazioti Deperet ; M. cretensis Simonelli.

The fragment from the loess of Kalouga attributed to " Cervus " verticornis by
Pavlow (1906 : 39) is not a Megaceros.

Megaceros verticornis, to be described in detail below, is the best known species.
It is nearly as large as the Irish giant deer, but more primitive in the dentition.
The basal portion of the antlers shows a remarkable constancy both in the form
and in the position of the brow tine. An accessory basal tine is quite exceptional.
The upper portion of the antler is broadly palmated, with a faintly scalloped edge.

Megaceros aff. verticornis from Siissenborn (Soergel, 1927) differs from the above
species by the presence of tines on the anterior edge of the palmation (the posterior
portion is destroyed). This might be perhaps a merely individual character. The
dentitions of these species are nearly identical.

Megaceros aff. verticornis from the Red Crag at Trimley (Dawkins, 1887) is repre-
sented by an imperfect shed antler. This antler is very stout but does not differ
otherwise from those of M. verticornis from the Forest Bed. Dawkins identified
it with this species, and were it not for its older age I also should not hesitate to
do so. The fossilization shows that this specimen is contemporary with the Crag,
but there is no exact information on the horizon in which it was found.

Megaceros dawkinsi from the Forest Bed will also be described below. It is
smaller than M. verticornis ; its forehead is prominent and the pedicles are strongly
directed backwards. The antlers are palmated, stout, small, and very variable in

52 DEER OF WEYBOURN CRAG AND FOREST BED OF NORFOLK

shape. The brow tine is fluctuating. I interpret these characters as due to a
secondary reduction. The dentition is hypsodont but with primitive features.

Megaceros belgmndi from Montreuil near Paris, described and figured by Belgrand
(1869) ; M. aff. belgrandi from Taubach (Pohlig, 1892) ; M. aff . belgrandi from the
high terrace of the Neckar near Laufen, Schwaben (Dietrich, 1909); and M. aff.
belgrandi from Tiraspol (partly determined as M. belgrandi, partly as M. euryceros
by Pavlow, 1906, pi. i, figs. 4, 5) are all very imperfect. They might perhaps be
identical with M. dawkinsi. According to Dietrich the specimen from Laufen is
of Upper Pleistocene age ; the others come from the Middle Pleistocene.

Megaceros mosbachensis from Mosbach (Soergel, 1927) agrees with M. verticornis
in the characters of the skull but shows some divergence in the basal portion of the
antlers. The peculiar upper brow tine is present only on the right side with an
accessory tine below it; on the left side there is only this accessory tine. The
teeth are massive and very broad. This species was based on a single specimen.

Megaceros solilhacus from the Middle Pleistocene of Solilhac, Haute Loire (Robert,
1829) is a very large species but with relatively small antlers. The palmation is
narrow, the brow tine is more or less reduced. A very fine antler has been figured
by Moullade (1886) and by Freudenberg (1914) under the name Cervus dama priscus.
The dentition is advanced, and the hyperostosis of the lower jaw is poorly developed.
This species inhabited the highlands of Central France during the Riss Glaciation.
I shall give soon a new description of it.

Megaceros algericus from the Middle and Upper Palaeolithic of Algeria and the
Mousterian of Morocco (Arambourg, 1938) clearly shows the skull characters of
this group. It reproduces also some features of M. pachyosteus, namely the extreme
degree of hyperostosis, the small size and the shortening of the muzzle. This is
well seen in a fine skull in the Palaeontological Museum in Paris, not yet described.

Megaceros pliotarandoides from the Upper Pliocene or Lower Pleistocene of
Cortiglione Monferrato, N. Italy (De Alessandri, 1903) is based on an incomplete
shed antler. It shows in its basal portion the characters of the group of M. verti-
cornis but is very unsatisfactory and its affinities are uncertain.

Megaceros cazioti from Corsica and M. cretensis Simonelli are dwarf island forms.
A more detailed discussion of these species will be given in another paper.

There are in addition some remains whose affinities cannot be established :
Megaceros dupuisi from the Lower (or Middle ?) Pleistocene of Rosieres, France
(De Grossouvre & Stehlin, 1912), is represented by some lower jaws and very
imperfect fragments of antlers. The teeth are identical with those I tentatively
identify with M. savini.

Megaceros sp. A large metatarsal from Leffe, N. Italy (Stehlin, 1930) seems the
only unquestionable evidence of the occurrence of this genus in the Villafranchian
of Italy.

Megaceros sp. A lower jaw from the Red Crag, in the Museum at Ipswich, not
yet described. The teeth are broken off and only a part of Mx is preserved. The
length of the tooth row is as in M. verticornis and the horizontal ramus is very thick :
its height below M2 is 50 mm., its breadth 40 mm. The fossilization shows that this
specimen was derived presumably from the lower part of the Red Crag.

DEER OF WEYBOURN CRAG AND FOREST BED OF NORFOLK 53

Megaceros sp. from St. Prest is possibly identical with M. dupuisi (Stehlin, 1912 :
206).

Megaceros cf. dupuisi from Csarnota and Piispokfurdo (Schaub, 1932) is also very
poorly represented.

A few giant deer have been described also from Switzerland (see Hescheler &
Kuhn, 1948: 190-192). One of them, from the Wengimoos near Berne, is remark-
able for the shortness of its neck, and is supposed to be post-glacial.

Several imperfect remains of giant deer have also been made known from Middle
Pleistocene deposits of England. They all belong to more primitive species than
M. giganteus.

According to Stehlin (1932 : 143) Cervus martialis Gervais (1859 : Z44> P^ 2I«
fig. 1-8) is also a Megaceros. I do not feel certain of this. The lower jaw (Gervais'
fig. i) is certainly of a giant deer, but the antlers seem rather to belong to a relative
of Euctenoceros sedgwicki.

Megaceros messinae Pohlig (1909) from Sicily seems to be another dwarf island
form.

Megaceros verticornis (Dawkins)

(FigS. 23 A, B, 24-27, 28 M, N, 29 E, F, 30, 32 B, C.)

1872. Cervus verticornis Dawkins (pars), p. 405, fig. 2.

1882. Cervus verticornis Dawkins : Newton, p. 61.

1887. Cervus verticornis Dawkins : Dawkins (pars), pi. 5, figs. 1-3 ; pi. 6, figs, i, 2 ; pi. 7,

figs, i, 2.

1891. Cervus verticornis Dawkins : Gunn (pars), pi. 2, fig. 97.
1891. Cervus verticornis Dawkins : Newton, p. 32.
1899. Cervus belgrandi Lartet : Harmer, p. 97, pi. 21.

SYNONYMY. The definition of this species has been discussed by Newton.
Dawkins, in his original description, included in it also an antler on which Newton
later based his Cervus dawkinsi. Newton chose as the type of C. verticornis the
basal fragment of an antler from Pakefield, figured by Dawkins (1872, fig. 2).

In his later description (1887) Dawkins again included in C. verticornis some
remains of other species. On p. 23 he recorded seven specimens with a double
basal tine; exact reference was made only to a specimen in the British Museum,
registered B.M. (G.D.), 33471. This register number includes a small sample of
very imperfect fragments, none of which belongs to Megaceros verticornis. A double
basal tine is quite exceptional in this species: I do not know the other specimens
quoted by Dawkins, but as yet I have seen only a fragment of a young individual
from Trimingham with a well-developed accessory basal tine (Fig. 26). This
specimen was found after the publication of Dawkins' works.

The antler from the Red Crag at Trimley figured by Dawkins (1887, pi. 6, fig. 3)
is imperfect and cannot, since it is older, be identified confidently with M. verticornis.
The fragment from Kessingland figured by Gunn (pi. 7, fig. 2) is doubtful ; it does
not seem to belong to M. verticornis.

OCCURRENCE. This is the commonest species of Cervidae in the Forest Bed and
has been recorded from nearly all the fossiliferous localities : Kessingland, Pakefield,

54 DEER OF WEYBOURN CRAG AND FOREST BED OF NORFOLK

Mundesley, Overstrand, Happisburgh, Trimingham, Hopton, Sidestrand, Bacton,
Cromer; it is common also in the Upper Freshwater Bed at West Runton, but is
not recorded at East Runton.

The remains of this species consist for the most part of basal fragments of the
antlers, either naturally shed or with fragments of the frontals. There are in
addition two complete brain cases, one in the Norwich Museum (Figs. 24, 25), the
second in the British Museum (Fig. 27). The latter was found in connection with
the atlas, the axis and the antlers, nearly perfect, and has been described by Harmer
under the name Cervus belgrandi, of which Cervus verticornis was incorrectly supposed
to be a synonym.

A shed antler, in the Museum of the Geological Survey, figured by Dawkins
(1887, pi. 5, fig. 2), is broken at the beginning of the palmation.

The correlation between skulls and teeth is based on the size, on their occurrence
together in the Upper Freshwater Bed at West Runton, where no other species of
Megaceros has as yet been found, and on comparison with M. aff. verticornis from
Siissenborn. The dentition is represented by some lower jaws and maxillae.

Some limb bones may also possibly belong to this species.

DESCRIPTION. Skull: There is very little to add to Harmer's description of the
skull from Pakefield. Harmer pointed out its large size and heavy form, its flat
forehead and its divergent and widely spaced pedicles. The supraorbital foramina
are very large and round. Two very faint ridges, starting from the coronal suture,
run along the inner margins of the pedicles and vanish on the forehead. In other
specimens these ridges are better developed (Fig. 25). Harmer also pointed out
the lack of a transverse swelling of the frontal between the pedicles.

The other remains of skulls display little individual variations. The pedicles are
generally widely spaced, but in a frontal from Pakefield (Figs. 23 A, 25 D) they are
set near each other and less divergent than in the average specimens. The frontal
is still flat.

Antlers: The characters of the lower portion of the antlers are remarkably con-
stant. The brow tine arises at some distance above the burr, from the upper portion
of the beam, and is strongly bent dowwards. It is rounded in cross-section and
very long in a skull from Kessingland (Dawkins, 1887, pi. 7, fig. 2), the only specimen
where it is complete. In many specimens there is a more or less marked knob
below it, on the anterior side of the beam; this may be present on both sides or
on one side only and is very variable. In a frontal from Trimingham, very much
worn, it is very strong on the left side and seems to be completely lacking on the
right side, but only in a young antler (Fig. 26), also from Trimingham, has it developed
into an accessory lower tine. Dawkins' statement that an accessory lower tine
occurs in seven other specimens has not been fully checked, but is based partly
at least on incorrect identifications.

The beam is long and bears an anterior and a posterior tine, ovoidal in cross-
section. The position of the anterior tine is somewhat variable (see Figs. 25 c, 27,
and the figures published by Dawkins, 1887). An additional anterior tine has been
recorded only in one instance (Dawkins, 1887, pi. 5, fig. 3). Above the posterior
tine the beam is twisted upwards and becomes flattened, and after a short distance

DEER OF WEYBOURN CRAG AND FOREST BED OF NORFOLK 55

56 DEER OF WEYBOURN CRAG AND FOREST BED OF NORFOLK

FIG. 25. Megaceros verticornis, skulls. A. Inferior aspect, from Kessingland. Norwich
Museum 364. (See also figs. 23 & 24.) B. Posterior aspect, same specimen. J
natural size. c. Anterior aspect. West Runton, Upper Freshwater Bed. B.M.
(G.D.), M. 6099. f natural size. D, Anterior aspect. Pakefield, Norwich Museum
365. $ natural size.

DEER OF WEYBOURN CRAG AND FOREST BED OF NORFOLK 57

it suddenly expands into a palmation. This palmation is preserved only in Harmer's
specimen, so that we have no information on its variations. It is very broad and
thin and of a quite unusual pattern : its margin does not give origin to long tines as
in most of the giant deer, but is scalloped and gently bent inwards. The anterior
edge was destroyed on both sides. The plane of the palmation is nearly vertical

FIG. 26. Megaceros verticornis, young specimen with an abnormal lower tine, Trimingham.
B.M. (G.D.), Savin 713. £ natural size.

FIG. 27. Megaceros verticornis, reconstruction of the specimen described by Harmer (1899),
Pakefield. B.M. (G.D.), M. 11352. Tx¥ natural size.

and set at 45° with the median plane of the body. The total span in the reconstructed
specimen is 228 cm. Harmer also pointed out the rectilinear course of the blood
vessels.

Harmer's specimen is rather large, but the lectotype and the skull from the
Upper Freshwater Bed of West Runton (Fig. 25 c) slightly exceed it. The antler
figured by Dawkins, 1887, pi. 5, fig. 2, the most complete beyond Harmer's speci-
men, is on the contrary distinctly smaller.

58 DEER OF WEYBOURN CRAG AND FOREST BED OF NORFOLK

Dentition: The remains consist of a maxilla and two lower jaws from the Upper
Freshwater Bed at West Runt on, a maxilla from Trimingham and some imperfect
lower jaws from Pakefield, in the British Museum; a maxilla from Kessingland
and probably two lower jaws, from Copt on and Kessingland, in the Norwich
Museum.

M

FIG. 28. Cross-sections of lower jaws, below hinder lobe of M2. External side to the
left, f natural size. A-I. Megaceros dawkinsi. A. Mundesley, B.M. (G.D.), M. 6335
(young). B. Overstrand, B.M. (G.D.), Savin 533. c. Sidestrand, B.M. (G.D.),
M. 6224. D. Overstrand, B.M. (G.D.), M. 6209. E. East Runton, B.M. (G.D.),
Savin 304. F. Mundesley, B.M. (G.D.), M. 6207 (old). G. Overstrand, B.M. (G.D.),
Savin 526. H. Mundesley, B.M. (G.D.), Savin 339. i. Pakefield, B.M. (G.D.),
M. 6501. j. Cervid indet., Overstrand. B.M. (G.D.), Savin 421. K & L. Megaceros
savini ? K. Trimingham, B.M. (G.D.), M. 6220. L. Mundesley, B.M. (G.D.), Savin
1008 (old). M & N. Megaceros verticornis. M. Overstrand, B.M. (G.D.), Savin 1198.
N. West Runton, Upper Freshwater Bed. B.M. (G.D.), Savin 1069.

The hyperostosis of the lower jaw is moderate (Fig. 28). The teeth are relatively
small and narrow. The basal columns are moderately developed, and in the maxilla
from West Runton the upper premolars show a very faint trace of a cingulum. P4 is
somewhat intermediate between the primitive and the advanced condition. The
enamel is moderately thick, the height of the crown also moderate.

DEER OF WEYBOURN CRAG AND FOREST BED OF NORFOLK 59

B

D

FIG. 29. Megaceros, lower dentition, ^f natural size. A. M. dawkinsi, Overstrand. B.M.
(G.D.), Savin 533. B. M. dawkinsi, Mundesley. B.M. (G.D.), M. 6231 (young),
c. M. savini ? Trimingham, B.M. (G.D.), M. 6207. D. M. savini ? Trimingham,
B.M. (G.D.), Savin 439. E. M. vevticornis, West Runton, Upper Freshwater Bed.
B.M. (G.D.), Savin 1069. F. M. verticornis, West Runton, Upper Freshwater Bed.
B.M. (G.D.), Savin 1265.

60 DEER OF WEYBOURN CRAG AND FOREST BED OF NORFOLK

VERTEBRAE. Atlas and axis were found in connection with Harmer's specimen.
They have already been described by Harmer and are slightly smaller than the
corresponding bones of Megaceros giganteus, but very similar in shape.

COMPARISONS. The giant deer from Siissenborn, figured by Soergel (1927, pi. 17,
fig. 2 ; pi. 18, figs. 2, 4, 7), is closely related to M. verticornis. The skull, the denti-
tion and the lower portion of the antlers are identical. The upper portion of the
antlers in the specimen from Siissenborn is flattened and gives off two large tines
from the anterior edge ; the posterior portion is destroyed. In Harmer's specimen
from the Forest Bed the anterior edge of the palmation is missing on both sides,
but what remains of it on the right side is very thin (15-20 mm.) and is not likely
to have developed large tines, which moreover would be in contrast with the general
pattern of the palmation.

The systematic value of these differences is not known, as we have no idea of the
range of individual variation of these forms, but no doubt they are very closely
allied.

FIG. 30. Megaceros verticornis, lower molars, Pakefield. B.M. (G.D.),
M. 6223. Natural size.

Megaceros mosbachensis, of which only a brain case with the lower portions of the
antlers and some fragments of the dentition are known, also closely resembles M.
verticornis in the characters of the skull, but differs in the antlers and teeth (see
above).

The affinities between M. verticornis and M . dawkinsi are also very close and will
be discussed later.

The lower jaw of M. dupuisi differs from that of M. verticornis by its smaller size
and the more primitive form of P4.

The affinities between M. verticornis and M. giganteus have been discussed by
Harmer and by Soergel. It has already been shown that although these two species
are related they belong to different lineages.

MEASUREMENTS OF THE TEETH : —

Upper tooth row :

Total length
Breadth of M*

B.M. (G.D.), Savin 1156

W. Runton,
U. Freshw. Bed

124
22

6i

B.M. (G.D.), B.M. (G.D.),

Savin 1069 Savin 1265

Lower tooth row : W. Runton, U. Freshw. Bed

Total length . 146 . 152

Breadth of M2 . 17 . 18

Megaceros dawkinsi (Newton)

(FigS. 23 C, 28 A-I, 29 A, B, 31 A, 32A, 33-36.)

1872. Cervus verticornis Dawkins (pars), p. 405, fig. i.

1882. Cervus dawkinsi Newton, p. 54.

1882. Cervus fitchii Gunn MS. : Newton, p. 56.

1882. Cervus gunni Dawkins MS. : Newton, p. 57.

1887. Cervus dawkinsi Newton : Dawkins, p. 7, pi. 2, figs, i, 2, pi. 3, fig. 3.

1891. Alces ? : Gunn, pi. 5, fig. 6.

1891. Cervus fitchii Gunn, pi. 6, fig. i.

1891. Cervus dawkinsi Newton : Newton, p. 26.

1891. Cervus fitchii Gunn : Newton, p. 28.

SYNONYMY. This species was founded by Newton on a young antler, formerly
attributed to Cervus verticornis by Dawkins. Other fragments determined as
Cervus fitchii and Cervus gunni by Newton belong^to^the same species, as already
pointed out by Dawkins. Newton (1891) rejected this view, but in my opinion
he overestimated differences due to individual variation.

OCCURRENCE. This species is nearly as abundant and widespread as Megaceros
verticornis and has been recorded from Bacton, Cromer, Trimingham, Sidestrand,
Mundesley, Overstrand, Pakefield, Walcot and East Runton (one specimen, much
rolled). A specimen in the Savin collection was found at Weybourn and was
supposed to come from the Weybourn Crag; its occurrence has been discussed in
the previous section. M. dawkinsi has not been recorded from the Upper Fresh-
water Bed.

The specimens consist of several portions of antlers and fragments of the f rentals ;
a specimen from the Walcot gap, in the museum at Norwich, bears also a part of
the left parietal and temporal bones, but no complete brain cases have been found.
The correlation between antlers and teeth is based on the size. The dentition is
represented by several lower jaws and few fragments of the maxillae.

DESCRIPTION. The skull is distinctly smaller than that of Megaceros verticornis,
but is represented by rather unsatisfactory specimens, so that exact measurements
are possible only for the frontals. Its most striking feature is given by the convex
form of the forehead and by the backward inclination of the pedicles. In the adult
the pedicles are short, very massive and strongly divergent. Two shallow ridges
on their inner borders, as in M. verticornis, are always distinct. The skull roof
of the specimen reproduced on Fig. 33 A is pierced by numerous small openings;
its inner surface in this region has been filled with plaster. Similar openings have
been described also by Soergel (1927) in the giant deer from Mosbach and Siissen-
born ; they were interpreted as canals for blood-vessels.

62 DEER OF WEYBOURN CRAG AND FOREST BED OF NORFOLK

The antlers, of which the upper portion is unknown, are stout, palmated, and very
variable. The beam is short and rounded. Its base, unlike that of M. verticornis,
does not grow in the same direction as the pedicle but is stretched outwards and
backwards, and is thinner than the pedicle. This is well seen also in a young speci-
men reproduced on Fig. 33 c. The brow tine and the accessory lower tine are very
variable, but are always more or less reduced. The type affords the only instance
of a well-developed lower accessory tine. Very often both these tines have dis-
appeared. The upper tines are moderately flattened and gently twisted upwards.

FIG. 31. Megaceros, lower jaws. £ natural size. A. M. dawkinsi, Overstrand. B.M.
(G.D.), Savin 526. B. M. savini ? Trimingham, B.M. (G.D.), Savin 439. c. M.
savini ? Mundesley, B.M. (G.D.), Savin 1008.

There may be one or two anterior tines, and a posterior tine is not always present.
The upper portion is more or less abruptly twisted upwards, sometimes resembling
features of M. verticornis, sometimes as in Fig. 36, rather recalling M . giganteus.
In the specimen shown on Fig. 23 c and 33 A the branching of the right antler takes
place in two different planes: this is probably due to an unusual shortening of the
beam between the anterior tine and the beginning of the palmation.

The teeth I refer to this species consist of several lower jaws, a fragment of a
maxilla and some isolated molars. The dental characters recall those of M. verti-
cornis; the teeth are however smaller, narrower, very hypsodont, and have a thick
enamel. The lower molars bear strong basal columns and anterior folds. P4 is

DEER OF WEYBOURN CRAG AND FOREST BED OF NORFOLK 63

always very primitive and is characterized by the poor development of its hinder
portion ; the latter is bordered by a small burr near the base of its outer wall.

The hyperostosis of the lower jaw is well marked (Fig. 28).

AFFINITIES. Dawkins (1887) pointed out the general resemblance of the antlers
with those of M. giganteus, but concluded that the affinities between these two
species are not very close.

As he limited his observations to the antlers, he failed to perceive the affinities
between M. dawkinsi and M. verticornis, which moreover was imperfectly known
until the discovery of Harmer's specimen, ten years later.

FIG. 32. Megaceros, upper dentition. Natural size. A. M. dawkinsi, Trimingham,
B.M. (G.D.), Savin 7. B. M. verticornis ? Trimingham, B.M. (G.D.), Savin 942.
c. M. verticornisy West Runton, Upper Freshwater Bed. B.M. (G.D.), Savin 1156.

The characters of the skull leave little doubt that Megaceros dawkinsi belongs to
the group of M. verticornis. The antlers support this conclusion : it has been shown
that a small brow tine, similar to that of M. verticornis, is not uncommon. This
never occurs in the group of M. giganteus.

However, Megaceros dawkinsi, in spite of its small size, is not primitive, and does
not belong to the same lineage as M. verticornis. The dentition displays simul-
taneously primitive and advanced features; the lower pre-molars are perhaps of
the most primitive pattern yet known, but both the hypsodonty and the thickness
of the enamel are no doubt advanced features. There is no hint of the broadening

64 DEER OF WEYBOURN CRAG AND FOREST BED OF NORFOLK

DEER OF WEYBOURN CRAG AND FOREST BED OF NORFOLK 65

GEOL. II, I.

66

DEER OF WEYBOURN CRAG AND FOREST BED OF NORFOLK

of the molars, undergone by Megaceros giganteus and M. mosbachensis. Nor can
the antlers be considered primitive. The pedicles are disproportionately strong for
their size; moreover, in cross-section the beam is thinner than the pedicle and is

FIG. 35. Megaceros dawkinsi, left antler, seen obliquely from the outer side. Trimingham,
B.M. (G.D.), Savin 1820. £ natural size.

FIG. 36. Megaceros dawkinsi, reconstructions of the antlers based on the specimens of

Figs. 33 & 34. £ natural size.

DEER OF WEYBOURN CRAG AND FOREST BED OF NORFOLK 67

stretched backwards — two common features in the deer with reduced antlers. The
unusually high instability of the tines is also a character peculiar to regressed
antlers.

Whether this regression of M . dawkinsi was due to inner factors or to influence
of the environment can hardly be stated. The characters of the dentition suggest
habits of life in open, grassy lands, rather than in woodlands; this is a rather
unusual habitat for deer.

Megaceros belgrandi and several related forms from Central and Eastern Europe,
listed above, are closely similar to M. dawkinsi and possibly identical with it, but
their remains are very imperfect.

A striking similarity with the antlers of M. dawkinsi is shown by an antler of
Cervus cazioti from the cave of Nonza in Corsica, figured by Deperet (1897^), which,
however, is less than half the size of the antlers of M. dawkinsi. Cervus cazioti
is no doubt a dwarf island form. Also its dentition recalls M. dawkinsi.

MEASUREMENTS : —

B.M. (G.D.) M. 6302
Frontal Sidestrand

Minimal breadth beneath the pedicles . 164

B.M. (G.D.), B.M. (G.D.), B.M. (G.D.), B.M. (G.D.),

Savin 526 M. 6224 Savin 339 Savin 533

Lower jaws Overstrand Sidestrand Mundesley Overs trand

Total length of the tooth row . 128 . 134 . 125 . 123

Breadth of M2 .... 15 . 15 . 15 14, 5

Megaceros savini (Dawkins)

(FigS. 23 D, 28 K, L, 29 C, D, 37-40-)

1887. Cervus savini Dawkins (pars), p. n, pi. 3, fig. 3, 5 ?

1891. Cervus savini Dawkins : Gunn (pars), pi. 7, figs. 4, 5.

1891. Cervus sp. Gunn, pi. 4, fig. 101, pi. 6, fig. 2.

1891. Cervus savini Dawkins : Newton, p. 30.

SYNONYMY. Dawkins included in this species also an antler of Euctenoceros
ctenoides (1887, pi. 3, fig. 4) and a fragment of a young specimen whose identity is
uncertain (pi. 3, fig. 2), but Newton pointed out that these identifications are
incorrect. Gunn attributed to it a frontal (pi. 4, fig. 102), which might possibly
belong to a red deer.

OCCURRENCE. This species is fairly frequent and is represented by not less than
twenty antlers in the British Museum and at Norwich. Eleven of them were found
at Trimingham, the others at Sidestrand, Overstrand, Mundesley and Kessingland.
A frontal from Kessingland (Figs. 23 D, 37) and possibly another frontal from Pake-
field (Dawkins, 1887, pi. 3, fig. 5) represent all that is known of its skull.

The remains of dentition I tentatively refer to this species consist of three lower
jaws from Trimingham and Mundesley, in the British Museum.

68

DEER OF WEYBOURN CRAG AND FOREST BED OF NORFOLK

DESCRIPTION. The frontal, which indicates an animal of about the same size as
Megaceros verticornis, clearly shows the hollowed forehead of the group of Megaceros
giganteus. The identification of the frontal (Figs. 23 D, 37) is made possible by the
basal portion of the antler.

The antlers include specimens of all ages. The type (Dawkins, 1887, pi. 3, fig. 3)
is of medium size, but the largest specimens, represented by basal fragments (Fig.
38 A, and a more imperfect fragment in the Norwich Museum, from an unknown

FIG. 37. Megaceros savini, frontal, Kessingland. Norwich Museum 321.
(See also Fig. 23.) J natural size.

locality), attained the size of Megaceros verticornis and M. giganteus. The beam is
flattened and hollowed below and bears a prominent ridge on its anterior side.
The brow tine (Fig. 38 c, D, and Gunn, 1891, pi. 6, fig. 2) branches off close to
the burr; it is flattened, broadly expanded into a small palmation, and set hori-
zontally. It is very often broken off at the base, and I do not know any specimen
where it is complete. The section of its base is triangular, with the longer side
above. The antlers are widely divergent, as in M. giganteus and in M. verticornis
(Fig. 39). The anterior and posterior tines are much flattened; above the posterior
tine the beam is bent upwards, becomes still more flattened and branches into two
terminal tines, apparently of equal size.

The teeth I tentatively refer to this species are intermediate in size between those
of M. verticornis and those of M. dawkinsi. The distinction is not sharp, and in

DEER OF WEYBOURN CRAG AND FOREST BED OF NORFOLK 69

their extreme variations these species, especially M. verticornis and M. savini, may
perhaps overlap. The lower tooth row is identical with that of Megaceros dupuisi
(Stehlin, 1912). P4 is primitive. The teeth I have seen are rather worn, but they
do not seem to be hypsodont. The enamel is of medium thickness.

FIG. 38. Megaceros savini, antlers. J natural size. A & B. Trimingham, large speci-
men. B.M. (G.D.), Savin 1422. c & D. Overstrand, medium sized specimen. B.M.
(G.D.), Savin 1880. E. Trimingham, young specimen. B.M. (G.D.), Savin 1241.

Two imperfect maxillae from Trimingham might perhaps belong to this species.

MEASUREMENTS : —

B.M. (G.D.), M. 6220
Lower jaw : Trimingham

Total length . . . 135

Breadth of M, 16

70 DEER OF WEYBOURN CRAG AND FOREST BED OF NORFOLK

FIG. 39. Megaceros savini, reconstruction of the holotype, B.M. (G.D.), M. 6093, and of
the specimens of Fig. 38. ^ natural size.

DEER OF WEYBOURN CRAG AND FOREST BED OF NORFOLK 71

FIG. 40. Megaceros savini ? Very young specimen. Bacton, B.M. (G.D.), Savin 910.

\ natural size.

FIG. 41. Megaceros, limb bones. £ natural size. A. M. verticornis or savini. Right

radius. No locality, B.M. (G.D.), Savin 1462. B. M. verticornis or savini. Right

metacarpal, Trimingham. B.M. (G.D.), Savin 1104. c. M. dawkinsi ? Right

metacarpal, Mundesley. B.M. (G.D.), Savin 2024. D. M. verticornis or savini,
Left metapodial, Trimingham. B.M. (G.D.), Savin 1450.

72 DEER OF WEYBOURN CRAG AND FOREST BED OF NORFOLK

Megaceros incertae sedis
(Fig. 41 A-D.)

LIMB BONES. Fourteen complete limb bones may be attributed to the genus
Megaceros for their large size and heavy form. They are: two radii, three meta-
carpals and four metatarsals in the British Museum; a metacarpal and a tibia in
the Norwich Museum ; a radius, a metacarpal and a metatarsal in the Museum of the
Geological Survey. The metapodials are characterized by the broad form of their
distal articulations and are easily recognizable.

The larger bones are of the same size as Megaceros giganteus and may be attributed
to M. verticornis or to M. savini ; the smaller ones probably belong to M. dawkinsi,
or to females of the larger species.

MEASUREMENTS : —

Radius :

Total length
Proximal breadth
Distal breadth

B.M. (G.D.),

Savin 464.

Trimingham

34°
79

B.M. (G.D.),
Savin 1462.
(No locality)

349

82

73

G.S. 21688.
Kessingland

320

Metacarpal :

Total length
Proximal breadth
Distal breadth

B.M. (G.D.),
Savin 1104.
Trimingham

304

64
68

B.M. (G.D.),
Savin 376.
Sidestrand

293
56
60

B.M. (G.D.),
Savin 2024.
Mundesley

286
63

Total length
Proximal breadth
Distal breadth

G.S. TF/55.
Mundesley

271
58

Norwich Museum

531-

(No locality)

310
70

74

B.M. (G.D.),

Savin 1885.

Sidestrand

261

48

52

Tibia

Metatarsal :

Total length
Proximal breadth
Distal breadth

Total length .
Distal breadth

G.S. 21684.
Kessingland

348
57
65

Norwich Museum

526.
(No locality)

482
81
B.M. (G.D.),

M. 6491.
Trimingham

358
59
67

B.M. (G.D.),
Savin 1450.
Trimingham

345
55
64

DEER OF WEYBOURN CRAG AND FOREST BED OF NORFOLK 73

CERVIDAE incertae sedis
" Cervus " obscurus n. sp.

(Figs. 42-45-)

SYNTYPES. B.M. (G.D.), Savin 195, Mundesley (Fig. 42 A, B); B.M. (G.D.), M. 2321, Forest
Bed : no exact locality (Fig. 42 C-E) ; B.M. (G.D.), M. 6421, Overstrand (Fig. 43).

ADDITIONAL SPECIMENS. Two lower portions of shed antlers and three basal
fragments of antlers with the f rentals; two of these (Fig. 44 c, D) possibly belong
to the same individual.

OCCURRENCE. Bacton, Mundesley, Sidestrand and Overstrand. In addition, an
antler of doubtful attribution from Trimingham.

DIAGNOSIS. A cervid of large size characterized by a small lower basal tine
directed downwards, a large upper basal tine strongly twisted outwards, a massive
beam and an upper tine set far apart from the basal tines.

DESCRIPTION. This is a species of very unusual features. The size is large and
the forms are massive as in Euctenoceros dicranios and ctenoides. The forehead is
marked by a prominent median suture and two shallow ridges along the inner
borders of the pedicles, as in Megaceros verticornis and its relatives, but is narrower.
A small tine is given off close to the burr, at a wide angle with the beam, and is
inclined inwards and downwards. A second basal tine, much more robust, arises
at some distance from the burr and is strongly bent outwards. It is rather long
and is flattened at its end. The beam is massive, straight, and ovoidal in cross-
section, with the major axis set vertically. A third tine, directed upwards, is given
off at a very great distance from the burr. The grooves of the blood-vessels are
shallow.

The remaining parts of the frontals enable one to reconstruct the position of the
antlers. The beams diverge at about 45° from the median plane of the body.

The antler from Trimingham which I doubtfully identify with this species (Fig.
44 A, B) differs in the characters of the lower tines. The lower tine arises at some
distance from the burr; a second tine arises from the inner side, near its base.
This tine corresponds for its position to the small knob between the two basal tines
of the syntype of Fig. 42 c, D, E.

AFFINITIES. The systematic position of this species is quite uncertain. No
doubt it does not belong to the genus Cervus in the restricted sense denned in the
previous pages. The lower portion of the antlers recalls that of Megaceros verti-
cornis, but the characters of the upper portion of the antlers and of the forehead
rule out any relationship with Megaceros] they rather point to some affinity with
Euctenoceros, but this question is still open.

Specimens of Doubtful Attribution

LOWER JAWS. Two lower jaws from Overstrand and Trimingham (Figs. 28 j,
46, 47) might belong either to " Cervus " obscurus or to Euctenoceros sedgwicki. They
both belong to young but fully grown specimens. The teeth are massive and

74 DEER OF WEYBOURN CRAG AND FOREST BED OF NORFOLK

FIG. 42. " Cervus " obscurus, two of the syntypes. J natural size. A & B. Mundesley,
B.M. (G.D.), Savin 195. c, D & E. Forest Bed> B.M. (G.D.), M. 2321.

DEER OF WEYBOURN CRAG AND FOREST BED OF NORFOLK 75

FIG. 43. " Cervus " obscurus. Syntype, Overstrand. B.M. (G.D.), M. 6421.

natural size.

FIG. 44. A & B. " Cervus " cf. obscurus, Trimingham. B.M. (G.D.), M. 6400. c.
" Cervus " obscurus, right antler of young specimen, Mundesley. B.M. (G.D.),
M. 6307. D. " Cervus " obscurus, left antler of a young specimen, Mundesley. B.M.
(G.D.), M. 6315, probably both belonging to the same individual. J natural size.

76 DEER OF WEYBOURN CRAG AND FOREST BED OF NORFOLK

seem to be slightly more hypsodont than those from East Runton which I tentatively
refer to Euctenoceros ctenoides. P4 has a complete internal wall and the molars
bear a strong anterior ridge. The ramus is depressed and thick, and its cross-
section closely resembles that of the lower jaws of Euctenoceros from the Upper
Valdarno (Azzaroli, 1948, fig. n, no. 3). The jaw from Trimingham, which is
more complete, is distinguished by a large coronoidal process. The breadth of M2
is 17 mm.

FIG. 45.

" Cervus " obscurus, tentative reconstruction, based on the syntypes.
size. A. Anterior view. B. Dorsal view.

natural

BRAIN CASE. The identification of a brain case from Trimingham (B.M. (G.D.),
M. 6303) is uncertain. The shortness of the pedicles gives evidence that it belonged
to a fully grown specimen ; they are, however, too small for Euctenoceros ctenoides,
E. sedgwicki or " Cervus " obscurus. Moreover, the forehead is hollowed, and bears
no trace of the ridges which distinguish the latter species. Euctenoceros tetraceros
is not recorded at Trimingham, and the supraoccipital crest of the specimen in
question is much stronger than that of the skull roof from East Runton I tentatively
identify with E. tetraceros. The brain case from Trimingham might possibly belong
to Cervus cf. elaphus', however, it exceeds the size even of the largest specimens
from Val di Chiana (Azzaroli, 1948).

MEASUREMENTS : —

Occipital breadth . . . . . .156

Occipital height ...... 93

Minimal frontal breadth . . . . .152

Breadth of the brain case between the parietals . 104
Breadth of the condyles . . . , .89

DEER OF WEYBOURN CRAG AND FOREST BED OF NORFOLK 77

FIG. 46. Cervid indet., lower dentition. Trimingham, Norwich Museum 45. Natural size.

FIG. 47. Cervid indet., Trimingham, Norwich Museum 45. (See also
Fig. 46.) £ natural size.

78 DEER OF WEYBOURN CRAG AND FOREST BED OF NORFOLK

LIMB BONES. An account of the complete limb bones is given in the following
table of measurements; some of them have been reproduced in Fig. 48. The
metapodials are easily recognizable from those of Megaceros by the narrower form
of their distal articulation, but a more precise determination is difficult and can be
based only on the proportions. They might be attributed to Euctenoceros, to the
red deer, or to " Cervus " obscurus.

FIG. 48. Cervids indet., limb bones. J natural size. A. Left metatarsal, West Runton,
Upper Freshwater Bed. B.M. (G.D.), Savin 1643. B. Left metacarpal, Trimingham.
B.M. (G.D.), Savin 2168. c. Left metacarpal, Trimingham. B.M. (G.D.), M. 6473.
D. Right metatarsal, Overstrand. B.M. (G.D.), Savin 773. E. Right metacarpal,
East Runton. B.M. (G.D.), M. 6476.

Some bones from Trimingham and from West Runton (B.M. (G.D.), Savin 2168,
1424, 1643) are long and slender like those of Libralces gallicus, but their shape is
quite different and leaves little doubt that they belong to a cervine. Their identi-
fication is puzzling; Euctenoceros ctenoides and E. tetraceros can be excluded, and
an identity with the red deer seems extremely improbable, although only this

DEER OF WEYBOURN CRAG AND FOREST BED OF NORFOLK 79

among the five species quoted above has been found in the Upper Freshwater
Bed at West Runton. Euctenoceros sedgwicki was presumably as heavy an animal
as the largest representatives of its genus. An identification with " Cervus "
obscurus seems less improbable.

MEASUREMENTS : —

Radius :

Total length
Proximal breadth
Distal breadth .

B.M. (G.D.), 1105.
Trimingham

337
70

63

B.M. (G.D.),M. 6469.
Sidestrand

325

Metacarpal :

Total length
Proximal breadth
Distal breadth .

B.M. (G.D.),
Savin 2168.
Trimingham

321
5i
50

B.M. (G.D.),
Savin 1424.
Trimingham

316
53
53

61

B.M. (G.D.),

M. 6476.
East Runton

B.M. (G.D.),

M. 6473.
Trimingham

Femur :

Total length

Proximal breadth .....
Antero-posterior diameter of the distal epiphysis

. 267 283 . 306 291
•45 50 • 55 52
• 45 50 • 57 53

B.M. (G.D.),
Savin 1446.
Trirningham

360

103
85

B.M. (G.D.), B.M. (G.D.), B.M. (G.D.), B.M. (G.D.),

Metatarsal :
Total length
Proximal breadth
Distal breadth

Savin 1685.
Trimingham

Savin 1071.
Overstrand

Savin 773.
Overstrand

M. 6490.
Sidestrand

310

307

311

265

42

4i

45

39

—

47

52

—

B.M. (G.D.),

M. 6495.
East Runton

Total length .
Proximal breadth
Distal breadth

293
44
50

293

47

B.M. (G.D.),

Savin 1643.

West Runton,

U. Freshw. Bed

347
46

54

Cervid cf . Dama nestii nestii

(Fig. 49-)

1882. Cervus etueriarum ? Croizet & Jobert : Newton, p. 55.
? 1889. Cervus rectus Newton, p. 145, fig. i, la.
? 1891. Cervus rectus Newton : Newton, p. 30, pi. 4, fig. i.

SYNONYMY. Newton's Cervus rectus is based on a very young specimen, probably of this species,
as may be seen by comparison of the type with fig. i n° i, 2 in Azzarpli, 1948.

OCCURRENCE. Seven lower portions of antlers and a fragment of the upper
portion, from East Runton; two basal fragments from Sidestrand and one from

FIG. 49. Cervid cf. Dama nestii nestii, antlers. J natural size. A. Left antler, Mundes-
ley. B.M. (G.D.), Savin 1164. B. Left antler, East Runton. B.M. (G.D.), M. 6365.
c. Fragment of the upper portion of a left antler, East Runton. B.M. (G.D.),
M. 6389. D & E. Left antler, East Runton. B.M. (G.D.), M. 6568.

DEER OF WEYBOURN CRAG AND FOREST BED OF NO,RFOLK 81

Mundesley. Probably two isolated M3 and some imperfect metapodials from East
Runton.

DESCRIPTION AND COMPARISONS. This species is poorly represented. The antlers
correspond with those of Dama nestii nestii from the Upper Valdarno and Olivola
(Azzaroli, 1948) for the position and development of the brow tine and the regular
helicoidal torsion of the beam. Also the fragment of the upper portion agrees with
the terminal fork of this subspecies ; the curvature of the beam and the characters
of the fork give evidence that the bifurcation took place in a transverse plane.
If so, the antler had four tines.

The other species, with which this species can be compared are : Cervus philisi
from Seneze (Schaub, 1942), Cervus rhenanus from Tegelen (Hooijer, 1947) and
Cervus perolensis from the Auvergne (Bout & Azzaroli, 1953). Cervus philisi
differs in the stronger development of the brow tine, the straighter form of the beam,
and the characters of the upper portion of the antlers. Cervus rhenanus, whose
antlers are more imperfectly known, differs mainly in the higher position of the
brow tine, and Cervus perolensis, also imperfectly known, in the stronger development
of the brow tine. These three species are rather primitive and their relationships
are not yet clear.

Cervid cf. Dama clactoniana Falc.

(Fig. 50)

I include here several specimens of the size of a fallow deer. Their identification
is somewhat uncertain and they might possibly include more than one species.
Their age is also uncertain, but at least the specimens from the Upper Freshwater
Bed are certainly post-Villafranchian.

ANTLERS (Fig. 50). Three basal fragments from Bacton, Trimingham, and West
Runton (Upper Freshwater Bed). They are distinguished by a very stout basal
bifurcation and a strong curvature of the brow tine. The pedicles are small. The
antlers from Bacton and from West Runton are disproportionately large for their
pedicles and probably belong to old individuals.

TEETH. Several isolated teeth and a complete lower tooth row from the Upper
Freshwater Bed at West Runton; two imperfect lower jaws, from Overstrand and
Trimingham. The lower molars bear well-developed basal columns. The P4
from Overstrand is primitive, two P4 from West Runton are advanced. The total
length of the lower tooth row from West Runton is 100 mm.

LIMB BONES. Two perfect metacarpals, from Overstrand and Trimingham.
Fragments of about the same size occur in the Upper Freshwater Bed of West
Runton.

MEASUREMENTS : —

B.M. (G.D.), B.M. (G.D.),

M. 6475. Savin 1399.

Metacarpal Overstrand Trimingham

Total length . 232 . 235

Proximal breadth -35-35
Distal breadth . 35 . 35

GEOL. II, I. 6

82

DEER OF WEYBOURN CRAG AND FOREST BED OF NORFOLK

COMPARISONS. In the Villafranchian fauna only Da-ma nestii eurygonos (Azzaroli,
1948) can be compared with the above specimens. However, the specimens from
Bacton and West Runton are larger than this subspecies; moreover those from
West Runton are certainly younger, and this is probably true also for those from
Bacton and Trimingham. After the close of the Villafranchian, no deer of this size
is known until the Clacton stage.

FIG. 50. Cervid cf. Dama clactoniana, right antlers. J natural size. A. Abnormal
specimen, Bacton, very old ? B.M. (G.D.), Savin 1283. B. Abnormal specimen,
very old? West Runton, Upper Freshwater Bed. B.M. (G.D.), Savin 554. c.
Trimingham, B.M. (G.D.), M. 6367.

In this horizon we find in East Anglia Dama clactoniana (Bate, 1938, Oakley &
Leakey, 1938), with which the fragments described above may be identified.

Keilhack (1888) described a pair of antlers of a large fallow deer from Belzig,
near Berlin, which he supposed to belong to the Lower Pleistocene or even to the
preglacial. But the occurrence of Alces in the same deposit points to an Upper
Pleistocene age.

DEER OF WEYBOURN CRAG AND FOREST BED OF NORFOLK 83

Species incorrectly recorded from the Forest Bed
" Cervus " polignacus Robert

1882. Cervus polignacus Robert : Newton, p. 59.

1891. Cervus polignacus Robert : Gunn, pi. i, fig. 94, pi. 2, fig. 95.

1891. Cervus polignacus Robert : Newton, p. 29, pi. 4, fig. 12.

Newton, on Falconer's authority, referred to Cervus polignacus two imperfect
antlers characterized by a low brow tine, branching at a very wide angle from the
beam. These two specimens have been figured by Gunn and are now in the Norwich
Museum. Several more imperfect antlers in the same museum have been
referred to this species, and in the British Museum I have seen other fragments
of a similar type. One of them, of a rather small size, has been incorrectly
identified with Megaceros savini by Dawkins (1887, pi. 3, fig. 2). All these
fragments are very unsatisfactory and are susceptible of different interpretations.

The name Cervus Dama polignacus was given by Robert (1829) to two incom-
plete skeletons from Solilhac: one of them is identical with his Cervus solilhacus
(now Megaceros solilhacus} from the same locality. The second is a small red deer
with abnormal characters; Pomel (1853) named it Cervus roberti.

" Cervus " carnutorum Laugel

A fragment said to come from the Chillesford Clay at Aldeby, identified with
Cervus carnutorum and figured by Dawkins (1872), may either belong to a large
specimen of Euctenoceros falconeri, or to E. sedgwicki. Three specimens from the
Norwich Crag quoted by Newton (1891, p. 26, pi. 4, fig. 3) possibly belong to E.
falconeri. Other specimens from the Forest Bed determined as Cervus carnutorum
in the collections belong to E. sedgwicki.

The name carnutorum, like polignacus, should be dropped (Stehlin, 1912).

Alces alces L.
1891. Alces ? Gunn, pi. 6, fig. 4.

A much rolled antler of Alces alces, from an unknown locality, has been figured
by Gunn. Its fossilization is less advanced than that of the other specimens, and
no doubt it came from a younger horizon.

Megaceros giganteus Blumenbach
1882. ? Cervus megaceros Hart : Newton, p. 58.

Although this species was recorded from the Forest Bed by the earlier authors,
Newton excluded it.

An isolated P4 from Sidestrand, in the Savin collection, registered B.M. (G.D.),
Savin 1601, might possibly belong to this species, but its occurrence in the Forest
Bed can be excluded.

84 DEER OF WEYBOURN CRAG AND FOREST BED OF NORFOLK

Rangifer tarandus L.
1891. Cervus sp. Gunn, pi. 6, fig. 3.

A much-rolled fragment of an antler, found on the beach at Pakefield, belongs
to this species. Gunn questioned whether it belonged to the Forest Bed Fauna,
and it can be excluded.

CHRONOLOGY

The Characters of the so-called Forest Bed Fauna

The so-called Forest Bed Fauna is very abundant in species, and includes repre-
sentatives of faunas met with at different horizons in continental Europe. This
unusual assemblage has been explained in various ways. Whereas the older authors
favoured an Upper Pliocene age (Reid, 1890, with bibliography), or assumed that
this fauna was partially or totally derived (Dubois, 1905), more recent authors
(Osborn, 1922, followed by Zeuner, 1945) attributed the whole fauna to the early
Pleistocene (post-Villafranchian), and interpreted the archaic species as relics.
Zeuner distinguished a " Cromer Forest Bed," where these archaic representatives
are found, and a " Bacton Forest Bed " without archaic elements, which he thought
to be a little younger.

All these interpretations are untenable. A Pliocene age is ruled out by the
presence of species which immigrated into Europe after the close of Villafranchian
times. On the other hand, the older representatives, once attributed to the Pliocene
but actually of Upper Villafranchian age, do not constitute isolated relics; an entire
faunal assemblage characteristic of that period is present. Moreover, primitive
species occur together with their more advanced descendants, and Zeuner's " Cromer
Forest Bed " is richer in species than any other locality. The deer, according to
the present revision, include 15 species, and we cannot imagine that they lived at
the same time. A derived fauna can also be excluded, as the fossils consist for a
large part of large antlers of deer, jaws and limb-bones of elephants, all of them
heavy and brittle, but rather well preserved. This is true both for the older and
for the younger representatives of the fauna.

The alternative solution, namely, that the deposit includes several horizons,
has been discussed in a previous section (" Geology "), and affords the only satis-
factory explanation of the mixed character of the " Forest Bed Fauna." An attempt
will be made here to determine the stratigraphical range of the 15 species of deer.
It will be useful, however, first to illustrate the time scale we shall use for com-
parison.

Faunal changes during the Lower and Middle Pleistocene

The Pleistocene continental sequence is marked by widespread faunal changes,
consisting of extinctions of old forms, immigrations of new forms and evolution
of primitive species into more advanced ones in situ.

Following the resolution made by the International Geological Congress in London
(1948), we shall place the Calabrian, and its continental equivalent, the Villafran-
chian, at the base of the Pleistocene. However, it is now generally realized that

DEER OF WEYBOURN CRAG AND FOREST BED OF NORFOLK 85

the so-called main Villaf ranchian fauna does not constitute such a uniform assemb-
lage as former authors assumed, and that it ranges over a relatively long lapse of
time.

The well known faunas of the pumiceous sands and conglomerates of the Mon-
tagne de Perrier in Auvergne (Ravin des Etouaires, Roca Neyra, Pardines) are older
than any of the other faunas hitherto attributed to the Villafranchian, and their
inclusion in this stage is misleading. These faunas are broadly contemporaneous
with those of the Roussillon (Deperet, 1897) and of the first lacustrine phase of the
Upper Valdarno (lignites with Tapirus arvernensis and laminated clays with plants ;
see Merla, 1949 : 51-57). In the Montagne de Perrier two horizons can be distin-
guished (Jung, 1946), but according to Prof. Bout of Le Puy (private information)
there is no break in the sequence and the series seems to range over a short interval
of time. In the three localities recorded above there is no trace of the cold climatic
phase that marks the beginning of the Pleistocene.

The fauna of the immediately following cold phase is perhaps represented in the
solifluction deposits of Vialette, in the highlands of southern France (see Bout &
Cailleux, 1951). In my opinion this cold phase might correspond to the Giinz
glaciation and marks the beginning of the Pleistocene.

In the Upper Valdarno basin the first lacustrine phase is overlain uncomformably
by a second lacustrine series (Merla, 1949), and to this only is the term Villafranchian
generally applied by Italian geologists. Its lower portion, not exposed, has been
explored by borings and has yielded a flora that seems to indicate a cold climate.
In the exposed section probably two horizons can be distinguished palaeontologi-
cally, a lower horizon with Mastodon arvernensis and an upper horizon with Elephas
meridionalis, although these two species may have lived together for a short period
(Merla, 1949; Azzaroli, 1950).

The fauna of Olivola (Azzaroli, 1950) is contemporary with the lower section with
Mastodon, that of Tegelen (Hooijer, 1947) with the upper section with Elephas.
The latter, according to Woldstedt (1950, " Tegelenschichten ") is of an imme-
diately pre-Elster age, that is, it belongs to the Giinz-Mindel interglacial.

The end of this period is probably represented by the faunas of some solifluction
deposits of southern France (Bout & Cailleux, 1951 ; Bout & Azzaroli, 1953, and private
information from Prof. Bout). Seneze (Schaub, 1944) is perhaps the most cele-
brated of these deposits. Other localities are Sainzelles and the Creux de Peyrolles.
Perhaps also the loessic deposit of St. Vallier (Viret, 1948) should be placed here.
These faunas are decidedly younger than that of Vialette and may be correlated
with the Mindel glaciation, that marks therefore the close of the Villafranchian.

In England the Pleistocene starts with the Red Crag (Lagaaij, 1952). The mam-
malian fauna of this horizon is represented by poor fragments, among the most
interesting of which are Mastodon arvernensis, Elephas cf . planifrons, two species of
Megaceros, and Euctenoceros falconeri. This horizon presumably corresponds to
the unexposed section of the Villafranchian Valdarno series. The equivalent of
the highest section of the same series, and of the clays of Tegelen, is represented
in England by the Weybourn Crag of Norfolk (see also later).

86 DEER OF WEYBOURN CRAG AND FOREST BED OF NORFOLK

It is no easy task to state the fate of all the species of the main Villafranchian
fauna, as the relationships of many of them with the representatives of later faunas
are not yet clear. Sus strozzii, Euctenoceros ctenoides, E. dicranios, E. tetraceros,
Dama nestii, " Cervus " philisi and its allies, Meganthereon meganthereon, Brachy-
prosopus vireti, Gazellospira torticornis, Nemorhoedus meneghinii, Procamptoceras
brivatense, Nyctereutes megamastoides and Acinonyx pardinensis have left no des-
cendants in the later faunas. This is possibly true also for Leptobos', a doubtful
Leptobos, however, has been recorded from Siissenborn (Merla, 1949 : 49). The
destiny of the horses is less clear, as this group still needs revision (Stehlin, 1932-
33:51 ff.). The same difficulty arises for many carnivores; Epimachairodus
survived for some time, and the larger felids, canids and bears might perhaps have
evolved into more modern species (Stehlin, 1932-33, Schaub, 1949). The voles,
represented in the Villafranchian by the primitive Microtus, underwent a gradual
evolution which sets them among the most useful leading fossils for the Lower and
Middle Pleistocene (Hinton, 1926; Schreuder, 1936, 1943, 1950; Heller, 1933,
1939; Van der Vlerk & Florschutz, 1950). A similar evolution was carried out
by the elephants, although things are here less simple than they were thought at
first (Trevisan, 1953). Also Libralces seems to have evolved further (see section
" Palaeontology "). Dicerorhinus etruscus has been recorded also from post-
Villafranchian deposits, but is represented there by more advanced forms (Soergel,
1923). Hippopotamus is found in the Villafranchian and in later times; its history
may be one of successive immigrations and retreats (Stehlin, 1932-33). Macacus
and Castor also survived, but their remains do not indicate whether they underwent
any great change. Trogontherium boisvilletti seems to have survived unchanged in
France and the British Isles (Schreuder, 1951).

The following phase is marked by the immigration of new faunal elements from
Eastern Europe or from Asia: primitive red deer (Cervus acoronatus, C, benindei
and other poorly known forms), the roe, wild boars of the group of Sus scrofa (pro-
bably represented by a distinct variety; Soergel, 1923), Gulo gulo, Bos primigenius,
Bison priscus.

The voles evolved from Mimomys into primitive Arvicola (A. greeni, bactonensis,
mosbachensis}. A primitive Microtus has also been recorded from Mosbach. Lib-
ralces seems to have evolved from L. gallicus into L. reynoldsi, and the elephants
from Elephas meridionalis into various races of E. antiquus and E. trogontherii.
The group of Megaceros verticornis flourished in this period and possibly evolved
in situ. Probably also the rhinoceroses and many carnivores are descendants of
Villafranchian ancestors.

The best representatives of this period are the main faunas of Mosbach and Siis-
senborn, and the fauna of Mauer, in Germany; the fauna of Tiraspol in Bessarabia
(Pavlow, 1906); the Needian of the Netherlands (Van der Vlerk & Florschutz,
1950), and some fossiliferous sands in the Upper Valdarno, around the village of
Bucine (horizon 3 in Merla, 1949 : 51). These sands overlie comformably the Villa-
franchian, with the interposition of a series of barren gravels. In England the
equivalent of these deposits is represented by the estuarine section of the Forest
Bed series.

DEER OF WEYBOURN CRAG AND FOREST BED OF NORFOLK 87

The dating adopted here is in contradiction with some current views and needs
explanation.

Soergel (1928) assumed an early Mindel, if not even a pre-Mindel age for the
fauna of Mauer. This was deduced from the occurrence of five horizons of weathered
loess above the fossiliferous sands — a rather unsafe way of arguing, inasmuch as the
age of the deposit depends on the number of glacial phases we admit. Soergel
assumed two stages in the Wiirm (Weichsel) glaciation, two in the Riss (Saale)
and two in the Mindel (Elster) ; but recently Woldstedt (1950) described four phases
in the Weichsel glaciation and three in the Saale. These are sufficient to explain
the occurrence of even more than five horizons of weathered loess above deposits
of the great interglacial, and any evidence for attributing Mauer to the Mindel
glaciation falls short. On the other hand, positive evidence for attributing these
faunas to the Mindel-Riss interglacial is afforded by the Needian of the Netherlands,
which corresponds to the Poludina-beds representing the great interglacial in North-
ern Germany (Woldstedt, 1950).

In 1932 Solomon placed the Forest Bed series of Norfolk in the first interglacial
and assumed that the overlying glacial drift included representatives of the Mindel
(North Sea Drift), Riss (Great Eastern) and Wurm (Little Eastern) glaciations.
But some of the difficulties met with in identifying Solomon's North Sea Drift
with the Mindel glaciation and with the Norwich Brickearth of southern Norfolk
were stressed a year before by Boswell (1931), and were given again by Boswell
and by Hazzledine Warren in the discussion following the presentation of Solomon's
paper. Solomon's evidence of a Mindel-Riss horizon within the glacial drift, assumed
to be afforded by his " mid-glacial sands," is not conclusive, as these sands are
barren and form no continuous horizon. In the dating adopted here the second
(Mindel) glaciation falls between the Weybourn Crag and the Estuarine Bed. Its
deposits, represented in southern Norfolk by BoswelTs Norwich Brickearth, may
have been cut out in north Norfolk by the erosion at the base of the Estuarine Bed.

The distribution of the Bovidae seems to have been influenced by geographic
or climatic factors. The bison is recorded at Mauer, Mosbach and Siissenborn,
whereas the aurochs makes its appearance in Central Europe during the Riss glacia-
tion (probably in a mild interstadial). On the other hand this species is not un-
common in the sands near Bucine in the Upper Valdarno, with Cervus cf . ela-phus
and Elephas antiquus, which, from their position, cannot be much younger than the
Villafranchian, whereas the bison is recorded in Italy only from the Upper Pleisto-
cene. Possibly Bos immigrated into Italy along the southern slope of the Alps,
whereas for some unknown reason it avoided Central Europe, and Bison did the
reverse. In the gravels of Tiraspol, with a faunal assemblage characteristic of the
stage of Mauer and the upper sands of the Upper Valdarno, Bos and Bison occur
together. Unfortunately Bos is represented only by limb bones (Pavlow, 1906).

A later phase is marked by the appearance of more advanced species of Arvicola,
evolved in situ, and by a widespread occurrence of Microtus, probably immigrants ;
but there does not seem to be any well-marked break in the fauna. Dama clac-
toniana, recorded only from the British Isles, belongs to this period.

The incoming Riss (Saale) glaciation seems to have extinguished Dama clac-

88 DEER OF WEYBOURN CRAG AND FOREST BED OF NORFOLK

toniana and many species of the group of Megaceros verticornis. Elephas primi-
genius, or a related form, occurs for the first time at the beginning of this phase,
but became common only after a mild oscillation, well marked at Steinheim a.d.
Murr. In this mild interstadial the giant deer are represented here by Megaceros
antecedens (Berckhemer, 1941) and the red deer by Cervus elaphus angulatus
(Beninde, 1937). In the following cold phase the red deer is represented by a
modern form and Coelodonta antiquitatis makes its very first appearance as an
immigrant. The date of the immigration of the reindeer is less clear. Soergel
(1943) recorded it from Steinheim, Mosbach and Siissenborn, but I have been unable
to check the dating of these specimens. They are all said to belong to the tundra
group (R. arcticus}. Also Ovibos moschatus has been recorded from Siissenborn
(Soergel, 1941).

At the end of the Riss glaciation (late Drenthian) the voles are represented by
modern forms. A good guide fossil for the post-Rissian is Megaceros giganteus.

The Distribution of the Deer of the Norfolk Coast
and their value for Stratigraphy

As stated above, the deer of the Forest Bed series sensu lato include representatives
of different faunas. Two faunas are quite distinct, whereas the existence of a third
fauna is vaguely indicated by the deer, and is demonstrated conclusively only by
other mammals.

The earliest fauna belongs to the upper section of the Villafranchian. It can be
correlated with the horizon of Tegelen, Seneze, and the ferruginous sands of the
Upper Valdarno, and is represented by Libralces gallicus, Euctenoceros tetraceros,
E. ctenoides, cervid cf. Dama nestii nestii. This fauna occurs in the Weybourn
Crag.

The second fauna belongs to the second interglacial and corresponds stratigraphi-
cally to the classic faunas of Mosbach (main fauna), Mauer, Siissenborn (main
fauna) and Tiraspol. Its representatives are Libralces reynoldsi, Cervus cf. elaphus,
Megaceros verticornis, Capreolus capreolus. This fauna occurs in the Estuarine
Section of the Forest Bed.

The third fauna, corresponding to a later horizon of the same interglacial, is
probably indicated by Dama clactoniana, associated with Cervus cf . elaphus, Mega-
ceros verticornis, Capreolus capreolus. This fauna occurs, partly at least, in the
Upper Freshwater Bed. Possibly there is no true break between the two younger
faunas.

The distribution of these species is not uniform throughout the outcrop of the
Forest Bed series s.l. The Villafranchian species are restricted to the western
section of the outcrop; they are very common at East Runton, where they are
not accompanied by later fauna! elements, and at Sidestrand, associated with
younger species ; rare at West Runton, Overstrand and Mundesley. Some doubtful
and much-rolled specimens have been found also at Pakefield, at the eastern extreme
of the outcrop. The second fauna occurs in all the localities, with the exception
of East Runton, where only a few much-rolled specimens have been found.

DEER OF WEYBOURN CRAG AND FOREST BED OF NORFOLK 89

The poorly represented fallow deer, supposed to represent the third fauna, is
recorded from Bacton, Trimingham, and from the Upper Freshwater Bed at West
Runton. Other representatives of this fauna have been recorded by Hinton from
the Upper Freshwater Bed at Bacton and West Runton (see later), but this fauna
might be more widespread.

Many localities have therefore yielded a mixed fauna, whereas a few of them have
yielded more uniform faunas. Of particular interest are East Runton, with a pure
Villafranchian fauna, and the Upper Freshwater Bed of West Runton, which, also
on geological grounds, can be assumed to include only representatives of the "third"
fauna. This distribution may give a clue to an approximate dating of the remaining
species, not known or doubtful in other districts.

Libmlces minor is recorded only at East Runton and Sidestrand; it belongs
therefore to the Villafranchian fauna.

Libralces latifrons from Happisburgh (doubtful at Mundesley, Cromer and Walcot)
may belong either to the second or the third fauna.

Euctenoceros sedgwicki, from Bacton and Mundesley, probably belongs to the
second fauna. The presence of the large fallow deer and the absence of Libralces
reynoldsi at Bacton would rather suggest a later age, but the absence of E. sedg-
wicki in the very fossiliferous Upper Freshwater Bed of West Runton is, perhaps,
more significant. This species is possibly present also at St. Prest.

Megaceros dawkinsi and M. savini are probably of the same age. They do not
occur in the Upper Freshwater Bed of West Runton, nor at East Runton. M.
dawkinsi has been recorded also from the Weybourn Beds at Weybourn by Savin,
but the name of the horizon was possibly taken from the locality and it would be
unsafe to rely on this statement. This species, or a related form, seems to have been
widespread in continental Europe.

The dating of " Cervus " obscurus is more doubtful, but this species probably
belongs to the second interglacial.

The Evidence Afforded by Other Mammals on the Age of
the Forest Bed Series

The list of the other mammals from the Forest Bed s.l. also gives the impression
of a mixed fauna. We need only mention the occurrence of four species of elephants,
a fact not recorded from any other locality in Europe.

The elephants still need revision, and it would be unsafe to assume the alleged
Elephas primigenius as evidence of the existence of an early Rissian horizon in
the Forest Bed series. But the evidence afforded by the voles (Hinton, 1926;
19260.) is more conclusive, and demonstrates the existence of an horizon equivalent
to the Swanscombe gravels.

The " shelly crag " at East Runton yielded Villafranchian representatives
(Mimomys pliocaenicus, M. intermedius, M. savini} , whereas the Upper Freshwater
Bed at West Runton yielded a mixed fauna: Mimomys intermedius, M. savini,
M. maiori, Evotomys sp., Pitimys arvaloides, P. gregaloides, Microtus arvalinus,
M. nivalinus, M. nivaloides, M. ratticeppoides. This is certainly not evidence of

90 DEER OF WEYBOURN CRAG AND FOREST BED OF NORFOLK

the presence of different horizons in the Upper Freshwater Bed, as the older species
may easily have been washed in. The age of the deposit is indicated by the younger
species and corresponds to the early Drenthian (Van der Vlerk & Florschiitz, 1950).
The same species of Microtus have been found also in the middle gravels of Swans-
combe (Schreuder, 1950). A correlation between the Upper Freshwater Bed and
Swanscombe has been made also by Hinton (19260, 336-337). Shortly later Mochi
(1929 : 179-181) proposed a correlation of the Norfolk sequence with the earliest
glaciations that agrees substantially with that stressed in these pages and differs
only in minor details.

SUMMARY

GEOLOGY AND CHRONOLOGY. The mammal bearing horizons of the Norfolk
coast include three horizons separated by unconformities: the Weybourn Crag,
the Estuarine Bed or Forest Bed s. sir. and the Upper Freshwater Bed. The whole
series rests on a levelled surface of chalk and is capped by glacial drift. The so-
called Forest Bed Fauna is actually made of three successive faunas. The oldest
one, from the Weybourn Crag, is of upper Villafranchian age and is assumed to
correspond to the first (Gunz-Mindel) interglacial. Its representatives are: Euc-
tenoceros ctenoides, E. tetraceros, Libralces gallicus, L. minor, and a small species
perhaps identical with Dama nestii nestii. The second fauna, from the Estuarine
Bed, is contemporary with the main faunas of Mosbach, Siissenborn, Mauer, Tiraspol,
and the horizon of Neede, and is assumed to correspond to the second interglacial.
Its representatives are: Megaceros verticornis, M. dawkinsi, M. savini, Libralces
reynoldsi, Cervus cf. elaphus, Capreolus capreolus, and possibly Libralces latifrons,
Euctenoceros sedgwicki and " Cervus " obscurus. The third fauna, from the Upper
Freshwater Bed and possibly also from the highest section of the Estuarine Bed,
is contemporary with the gravels at Swanscombe and with the early Drenthian and
corresponds to a later period of the same interglacial. It is represented by a species
possibly identical with Dama clactoniana, together with Megaceros verticornis,
Cervus cf. elaphus and Capreolus capreolus. Conclusive evidence of its age is afforded
by small rodents.

PALAEONTOLOGY. Alcinae and Capreolinae were differentiated from the Cervinae
before the Upper Miocene. The Upper Miocene Cervinae belong to two quite dis-
tinct lineages. In eastern Europe they are represented by Damacerus bessarabiae
(= Cervocerus novorossiae} and D. variabilis (= Procervus variabilis), and may per-
haps have evolved into the Villafranchian Cervus ramosus. In China they are
represented by three-tined deer (incorrectly identified with Cervocerus novorossiae
by former authors), from which probably most of the Pleistocene and living Cervinae
took origin. The phyletic development of antlers is also discussed.

The deer of the Weybourn Crag and Forest Bed s.l. belong to the genera Libralces,
Capreolus, Cervus, Euctenoceros, Megaceros and possibly Dama; there is in addition
a species of Cervinae of uncertain affinities.

Libralces is a close ally of Cervalces. L. gallicus is smaller than living elks, but
L. reynoldsi, which is probably its descendant, is the largest deer hitherto known.
L. latifrons and L. minor are poorly represented.

DEER OF WEYBOURN CRAG AND FOREST BED OF NORFOLK 91

The roe also is poorly represented.

The earliest representatives of Cervus s. str. were more primitive than the living
European red deer; some of them were similar to living red deer of Central Asia.
The red deer of the Forest Bed also seem to be primitive, but their remains are
not satisfactory.

Euctenoceros is distinguished by peculiar features of the antlers. Its most primi-
tive representative is E. falconeri from the Red Crag. E. sedgwicki is possibly its
descendant. E. ctenoides (= E. teguliensis) and E. tetraceros do not differ from the
types from other Villafranchian localities.

Megaceros can be divided into two groups, based on characters of the skull and
antlers: the group of M. giganteus and the group of M. verticomis. M. savini
is a primitive species of the group of M. giganteus and its antlers are not palmated.
M. verticomis, the commonest species of deer in the Forest Bed, has large and widely
palmated antlers. M. dawkinsi belongs to the same group; it is smaller and has
secondarily reduced antlers, and a hypsodont dentition.

" Cervus " obscurus is a large species of quite unusual features and of unknown
affinities. Its remains are scanty.

Two much smaller species may possibly be identified with Dama nestii nestii
and with Dama clactoniana.

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PRESENTED

7 JUL 1953

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2 6 NOV 1953

A CONIFEROUS PETRIFIED
FOREST IN PATAGONIA

M. G. CALDER

BULLETIN OF
THE BRITISH MUSEUM (NATURAL HISTORY)

GEOLOGY Vol. 2 No. 2

LONDON : 1953

BULLETIN OF
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No. 2 (1949)- A Reconsideration of the Galley Hill Skeleton. K. P.

Oakley & M. F. Ashley Montagu ...... 55.

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A CONIFEROUS PETRIFIED FOREST IN

PATAGONIA

BY

MARY GORDON CALDER

(University of Manchester)

Pp. 97-138 ; Pis. 1-7 ; 7 Text-figures

BULLETIN OF
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GEOLOGY Vol. 2 No. 2

LONDON : 1953

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A CONIFEROUS PETRIFIED FOREST IN

PATAGONIA

By MARY G. CALDER

SYNOPSIS

The present paper contains the results of a detailed investigation of a large collection of
silicified coniferous remains from the remarkable petrified forest of Cerro Cuadrado, Patagonia.

Conclusive evidence of only two species (Araucaria mirabilis and Pararaucaria patagonica),
based on seed cones, has been found. Araucaria mirabilis belongs to section Bunya of the
genus Araucaria, and the affinities of Pararaucaria patagonica are probably with the Taxodiaceae :
both species are extinct. The associated wood, branches, twigs and seedlings probably belonged
to one or other of the species described. The age of the forest is uncertain, and is at present
placed somewhere between Middle Jurassic and Wealden.

INTRODUCTION

THE existence of petrified coniferous plant remains in regions west of the mouth
of the Rio Deseado in the province of Santa Cruz, Patagonia, appears to have been
first noted scientifically by Dr. A. Windhausen of the Argentine Geological Survey.
In 1919 he discovered silicified cones which were kept as curios in stores and farms
in the country south-west of the Gulf of San Jorge ; and in 1923 he found abundant
petrified material of conifers in situ in the region of the Cerro Alto (or Cerro Chato),
near the Estancia Belgrano, in a locality about S. lat. 47° 50' by W. long. 68° 40'
(Windhausen, 1924 : 203, footnote). His collection of silicified wood and cones
was passed on to Professor Gothan of Berlin, who gave a short description of them
(Gothan, 1925). Dr. Carlos Spegazzini had also (1924) briefly described silicified
twigs and cones, the latter similar to those found by Windhausen, which he had
received from various donors : these specimens were from the region of the Cerro
Madre y Higa, a small volcanic peak about 30 miles north-east of Windhausen's
locality (Spegazzini, 1924). Meantime, during 1923-24, a palaeontological expedi-
tion from the Field Museum of Natural History, Chicago, had been collecting fossil
mammals in Eastern Patagonia, and Dr. Elmer S. Riggs and others made an addi-
tional expedition to the region of the Cerro Madre y Higa, and its neighbouring peak
about 10 miles to the west, the Cerro Cuadrado, where they found " a considerable
number of fossil trees, some with stumps standing, others lying prone with broken
branches and cones scattered about them, revealing a forest of fossil Araucaria or
Brazilian Pines preserved on the site where it had grown " (Riggs, 1926 : 544 ;
Wieland, 1935 : 6).

These discoveries made it clear that in the region of the volcanic peaks of the
Cerro Cuadrado, Cerro Madre y Higa, and Cerro Alto, in a district about 10° south
of the modern forest of Araucaria araucana (Molina) (= A. imbricata Pav.)
in Chile and Argentina, and about 26° south of the forests of Araucaria angustifolia
(Bertol.) ( = A. brasiliana Rich.) in Southern Brazil, there existed the petrified

GEOL. II, 2 7

too A CONIFEROUS PETRIFIED FOREST IN PATAGONIA

remains of an extensive araucarian forest. The large Field Museum collection
made by Riggs and his co-workers was later reviewed by Wieland (1929, 1935) ;
his vividly-presented memoir, however, was without detailed anatomical studies
of critical features. A fuller account of one of the cones in the Field Museum,
with special reference to the embryo, was given by Darrow (1936). After this,
little further botanical investigation of the petrified forest was undertaken, subse-
quent accounts by Darrah (1939 : 222), Florin (1940 : 36), Arnold (1947 : 313) and
Andrews (1947 : 175) being recapitulations of earlier studies, though Florin's later
work (1944 : 513) gave a partly revised interpretation of the structure of one type
of cone. A non-botanical account of the petrified forest is presented in the memoir
by Wehrfeld (1935) of a journey through Santa Cruz. The geology of the regions
between the Rio Deseado and the Rio Chico where the forest is situated has also
been recently reviewed by Feruglio (1949 : 118 ; 1951 : 35), with special reference
to fossil floras of plant compressions found at other localities in this area.

Important new collections of the conifers of the petrified forest have, however,
been made by Dr. Franz Mansfeld in the region of the Cerro Alto whilst searching
for fossil vertebrates, and in 1936 the British Museum (Natural History) acquired
an extensive collection of some 600 specimens from him, some by presentation
and some by purchase. Similar material is also widely distributed in other museum
and departmental collections : a demonstration of some of it was given by W. T.
Gordon at the Geological Society of London on behalf of Dr. Mansfeld (Gordon,
1936). The collection in the Geology Department of the British Museum is a repre-
sentative one ; in it are included silicified woods, branches and twigs of various
sizes, seedlings, and cones, in varying states of preservation and offering remarkable
evidence on the nature of the conifers of the petrified forest. The present account
is based mainly on this material, though reference has also been made to specimens
in other collections : Professor Gothan has described some of this more recently-
collected material from specimens in Berlin and elsewhere (Gothan, 1950) and
reference is made to this description when the seedlings are discussed. A more
detailed study of these South American petrified conifers, and in particular, of
the cones, appears to be desirable, especially in view of recent developments in
studies of the taxonomy and morphology of the living Araucariaceae (White, 1947 ;
Wilde & Eames, 1948, 1952 ; Buchholz, 1949) and of the fossil Araucariaceae
(Kendall, 1948, 1949; Cookson & Duigan, 1951). In addition, the remarkable
extension and organization of knowledge of the taxonomy, morphology and distri-
bution of fossil conifers in general provided by the work of Florin (1931, 1940,
1938-45) give a more critical basis for comparative interpretation than was pre-
viously available.

LOCALITY AND AGE

The site of the petrified forest is in the neighbourhood of the small volcanic peaks
of the Cerro Alto (Chato), Cerro Cuadrado, and Cerro Madre y Higa, to the south
of the Rio Deseado in the region between lat. 47°-48° S., long. 68°-69° W. (cf.
Gothan, 1925, pi. 10 ; Wieland, 1935 : 6, text-fig, i). More recent accounts of the
area where the silicified plant remains are found have, however, shown that the

A CONIFEROUS PETRIFIED FOREST IN PATAGONIA 101

extent of the forest is considerably wider than suggested in earlier reports. Feruglio's
survey (1949 : 128, map opposite p. 134 ; 1951 : 69, 74) indicates a number of
additional localities where cones and branches occur, some 25-30 miles to the north-
west of the Cerro Alto : these are, Estancia Los Toldos, Canadon de las Cuevas,
and west of Cerro Botellon. According to Feruglio's maps, there are at least two
localities in the region south of the Rio Deseado called " Cerro Cuadrado " : one
lies between the Cerro Alto and the Cerro Madre y Higa in the area originally indi-
cated for the petrified forest, and another some 70 miles to the south-west near to
the Estancia La Guitarra. From the latter locality Feruglio has described a quite
separate mixed flora consisting of compressions including some araucarian remains
(1951 : 62-65, 69), to which reference will later be made. Feruglio's review of the
region of the Cerro Alto where the petrified araucarian remains are found (1949 :
128) does not include the vicinity of the Cerro Cuadrado, after which Wieland named
the forest, as one of the fossiliferous localities ; the Cerro Alto appears to be the
central locality of the forest, and also to be one which has produced very valuable
and representative collections, though no doubt much remains to be discovered
in other areas..

It is clear, from the various descriptions, that the forest was overwhelmed in
an outburst of volcanic activity, the plant remains being embedded in a rhyolitic
volcanic ash. Some are found lying weathered out on the surface, showing varying
degrees of erosion of surface features, and some of the tree stumps are still standing
as noted, for example, by Riggs (1926 : 544), and illustrated by Frenguelli (1941,
pi. 25, fig. i) ; Frenguelli's photograph shows widely scattered trunks, and not a
close-set forest formation. Numerous seedling stems, to which reference will later
be made, also tend to confirm the impression that the forest was preserved in situ,
and was overwhelmed whilst regeneration was in progress.

The exact geological age of the forest is still doubtful. Dr. Mansfeld suggested
that the rocks from which his specimens were collected were not older than Cretace-
ous, and that they might even be of Tertiary age ; these estimates appear to have
been based on the comparatively modern appearance of the plant structures, rather
than on geological data (Gordon, 1936). The horizon has been placed, in other
accounts of the petrified forest, as widely apart as mid-Triassic (Windhausen,
1924 : 203 ; 1931 : 201 ; Gothan, 1925 : 197 ; Wieland, 1929 : 60 ; 1935 : 8),
Cretaceous (Florin, 1940 : 36 ; 1944 : 513) and Eocene (Frenguelli, 1933, cf. Barrow,
1936 : 328, and Fossa-Mancini, 1941 : 68). Dr. A. F. Leanza, of the University Museum
of La Plata, Argentina, very kindly informed me (in lilt,, igih June, 1948) that
geologically it could then only be indicated that the forest belonged to some stage
in the Mesozoic. Windhausen dated the forest as Triassic in the belief that the
volcanic ashes which enclose the plant remains form part of an eruptive which,
in the geological correlations then in vogue, was comparable in age with the eruptive
cycle which in the province of Mendoza originated the porphyritic series found
beneath sediments containing a Thinnfeldia flora ; eruptive cycles are, however,
now known also to have occurred in Patagonia at later stages in the Mesozoic. Ferug-
lio has reviewed again, in some recent publications (1949 : 118 ; 1951 : 35) to which
Mr. W. N. Edwards very kindly drew my attention, the problem of the geologic

102 A CONIFEROUS PETRIFIED FOREST IN PATAGONIA

age of the forest. He describes a series of floras of plant remains in the form of
compressions from a number of outcrops of the complex of volcanic sediments
(the " porphyritic series of Bahia Laura ") in the region between the Rio Deseado
and the Rio Chico ; and compares them in detail with floras of mid- Jurassic to
Wealden age, with special reference to species of Hausmannia, Cladophlebis, Nils-
sonia, and Ptilophyllum. He concludes that the age of the volcanic series is in
large part between mid- Jurassic and Wealden ; and that the age of the petrified
forest around the Cerro Alto, which occurs in the same complex (although its exact
stratigraphic position in relation to the other floras has not been determined),
must lie somewhere within the same range (1949 : 131 ; 1951 : 74). Feruglio also
points out that a major unconformity separates this volcanic series from the Upper
Cretaceous ; and also that the original determination of a species of Estheria from
this series in this area as being of Rhaetic age is not now generally accepted. These
latter points further support his delimitation of the age on the basis of the floras.

PRESERVATION AND TECHNIQUE

The plant remains are silicified : the replacement of the organic matter has been
fairly complete, and the specimens have not been effectively sectioned by any
adaptation of the cellulose peel technique. Thin petrological slices have accordingly
been used (prepared by Mr. J. Fowler of Sheffield) where details of the tissues are
required. However, in some specimens or parts of specimens the silicification has
gone so far that little detail can be distinguished : in others the wall structure is
well denned, and such details as the nature of pitting in the xylem, and the structure
of the embryos in some of the cones, may be observed. The cut and polished
faces of the cones may be used quite effectively for general morphology in many
cases ; Wieland relied very largely on this means of examination in his description
of the types (Wieland, 1935), which was accordingly lacking in histological details
that have proved to be of considerable importance to the interpretation of the
general morphology and taxonomy. Cut and polished specimens form very pretty
curios, as the preservation is in rather light silica enhanced by a variety of chalce-
donic shades of colour. Darrow (1936 : 330) has already commented on the ten-
dency for the various tissues of the seeds to be preserved in distinctive shades of
quartz.

(i) FOLIATED AND DEFOLIATED TWIGS AND BRANCHES

Araucarites sanctaecrucis n. sp.

(PL i, figs. 4-6, 8-13)

1924. Araucarites ? Spegazzini, p. 133, text-fig. 4 (8).

1935. Proaraucaria mirabilis (Speg.) : Wieland, p. 27, pi. 13, fig. i.

DIAGNOSIS. Woody branches of araucarian habit, radially symmetrical, 5-25
mm. in diameter, occasionally with lateral axillary branches in one row or two oppo-
site rows. Evergreen : leaves or leaf-scars arranged in a spiral with angle of diver-
gence 3/8 or 5/13, the latter on the thicker branches. Leaves imbricate and more
or less appressed. Leaf rhomboidal, 8-14 mm. in length, 4-8 mm. in width at its

A CONIFEROUS PETRIFIED FOREST IN PATAGONIA 103

broadest region, tapering distally to a subacute, somewhat incurved apex, the mar-
gins converging at the apex at an angle of 6o°-8o°. Abaxial surface sometimes
showing parallel longitudinal ridges and grooves. Free part of leaf flat, about half
the length of the entire leaf, and seated on a flat leaf-base cushion, from which on
older branches it has been shed. Leaf-base cushions on older branches 20-30
mm. wide, 5-10 mm. high, with marked longitudinal fissures. Venation of leaves
and structure of epidermis not known. Secondary xylem, seldom preserved, with
uniseriate and contiguous bordered pits on radial walls of tracheids.

LECTOTYPE. The specimen figured by Spegazzini, 1924, fig. 4 (8).

HYPOTYPE. V. 30941. Geol. Dept., B.M. (N.H.). The specimen illustrated in
this paper (PI. I, figs. 8, 10, n).

LOCALITY AND HORIZON. Cerro Alto, Santa Cruz, Patagonia : Between mid-
Jurassic and Wealden.

DESCRIPTION. — Many of the detached branches show one side of the specimen
much weathered, with surface features nearly obliterated, and the other with
excellent preservation of external morphology (PI. i, figs. 8, 10, n). No doubt the
better preserved surface was that which lay downwards on the forest floor, whilst
the upper surface was subject to processes of weathering as the surrounding rhyo-
litic volcanic ash was worn away ; some specimens of leafy branches are still em-
bedded in the ash (PI. i, fig. i at L), but most are detached. PI. i, figs. 4, 9, 10, n
and 13 illustrate the habit of branches of varying age. PL i, fig. n shows an un-
branched stem probably four years old, while figs. 4 and 9 represent thinner branches,
probably penultimate branches of the shoot, and show axillary branches similar to
that (? i year old) in fig. 13, which tend to be arranged in two opposite rows or one
row on the parent branch. This condition is found in the penultimate branch sys-
tems of many conifers where dorsiventrality in respect of the ultimate branching
is common. The defoliated branch of lower order (PI. i, fig. 6) may represent the
broken apical region of a terminal main branch, with 3 laterals in an apparent whorl.

In the foliated shoots the leaves are arranged in spiral phyllotaxy with angle
of divergence 3/8 or 5/13, the latter on the thicker branches : Church (1904 : 99)
has noted such spirals of a higher order on the thicker leafy branches in the Recent
Araucaria excelsa. PI. i, fig. 10, t, shows the free lamina of the leaf, and at b the
leaf-base cushion, from the stem shown in PI. i, fig. n. This specimen demon-
strates that these plants were evergreen : in transverse section as seen in the cut
surface at the upper end (PI. i, fig. 8), it shows poorly preserved secondary xylem
with 3-4 rather ill-defined growth rings, probably annual. It also shows in some
patches of the poorly preserved secondary wood that the radial pitting of the
tracheids is uniseriate and contiguous as illustrated in PI. i, figs. 3, 7. The trans-
versely cut face (PI. i, fig. 8, t) also shows the free part of the leaf in sectional view,
demonstrating its flat, unkeeled nature. No veins are preserved in the leaf, so
that the broad-based leaf habit cannot be definitely associated with parallel venation :
nor can the longitudinal striation which is seen on the abaxial surface of the leaf
in some specimens (e.g., PI. i, fig. 5) be definitely associated with any anatomical
feature such as ribs of sclerenchyma. The furrows in such cases may represent the
position of lines of stomata, but it has not been possible to demonstrate these structur-

104 A CONIFEROUS PETRIFIED FOREST IN PATAGONIA

ally, either from sections or cuticles. The older defoliated branch seen in PI. i,
fig. 12, probably about 5 years old, shows tangentially-extended leaf-base cushions
without any clearly-defined scar of attachment of the free lamina of the leaf : it is
doubtful whether the longitudinal fissuring in such specimens is entirely a natural
condition following on the stretching of the bark with increase in secondary thicken-
ing, but it is a common feature in some living araucarians (cf . A . araucana, Seward &
Ford, 1906, pi. 23, fig. D).

In none of the specimens figured or described in previous accounts of the forest
is there any indication of foliated branches : those referred to by Spegazzini (1924)
and by Wieland (1935) as Araucarites ? and Proaraucaria mirabilis respectively
are similar to that figured in PI. I, fig. 12 of the present account, which has been
interpreted as an older branch where the free lamina of the leaf has been shed.
After comparison of the variation in morphology of branches of different age in, for
example, the living Araucana araucana (cf. Seward & Ford, 1906, pi. 23, A-G), there
would seem to be no reason for regarding these thicker defoliated branches as a
separate species in the absence of any further criteria of difference : the structure
of the secondary xylem is too infrequently and inadequately preserved to be used
as such. Various references to similar branches (Darrow, 1936 : 333, 337 — where
they are identified with Proaraucaria mirabilis, — and Arnold, 1947 : 314) have
compared them with the living South American species of Araucana, A. araucana
and A. angustifolia, which comprise the section Columbea Endlicher, emend. Wilde
& Eames. However, from the locality Meseta de Baquero, about 70 miles to the
south-west of the Cerro Alto petrified forest, Feruglio has recently described com-
pressions of leafy branches as a new species of Araucana, A. grandifolia (Feruglio,
1951 : 62, pi. 3, fig. 5), and he suggests that these might belong to the cone species
Proaraucaria mirabilis, from the petrified forest, whose vegetative organs, he says,
are not known (1949 : 137). He adduces no botanical reasons for this suggestion
of relationship, and it is presumably based on geographical propinquity. Comparison
of the habit of A . grandifolia with that of the leafy branches here described certainly
does not suggest specific identity between the sterile branches from the two localities,
for A . grandifolia has much larger leaves, long, rather narrow, pointed and lanceolate,
and of spreading habit. There are no cuticular studies of this species, and on the
evidence of the external morphology alone one might even hesitate to assign the
branches to the genus Araucaria.

Although the general habit of the foliated shoots from the petrified forest suggests
comparison with an araucarian type, they cannot be identified on the basis of their
external morphology with any living species of Araucaria. It is true that the flat
and rather broad laminae of the leaves more closely resemble those of the living
species of the sections Columbea, Bunya and Intermedia of the genus Araucaria than
they do those of section Eutacta (cf. Wilde & Eames, 1952 ; White, 1947) ; but the
combination of characters represented by their relatively small size and flat, some-
what scale-like habit, broad base and rhomboidal form and appressed and imbricate
arrangement cannot be matched in any known living species. Indeed, in the absence
of information on leaf venation and epidermal characters there is no sound justifica-
tion for their inclusion in the genus Araucaria ; and the only other existing genus

A CONIFEROUS PETRIFIED FOREST IN PATAGONIA 105

in which they might conveniently be placed is Araucarites. Araucarites has been
used to define fossil cones or shoots of araucarian habit, for various reasons not
identifiable with the genus Araucaria itself (Seward, 1919 : 256), — though Seward
pointed out (pp. 265-6) that the use of this generic name for sterile shoots is not in
accordance with sound principles. More recent detailed studies of " araucarian "
sterile shoots (Brachyphyllum, Pagiophyllum : Kendall, 1948, 1948^, 1949) have
demonstrated that fossil genera which may be distinguished from Araucaria, in
cuticular structure may have this habit ; though in the case of Brachyphyllum
mammillare (Kendall, 1949) it is almost certainly related to cone structure resembling
that of Araucaria. Where, as in the present material, it is not possible to make
cuticular investigations, or any definite assignation to fertile material, it is most
convenient meantime to retain the artificial genus Araucarites to include these
branches, which are certainly of araucarian habit. As will be noted in a later
section of this paper, the sterile branches are associated with seed cones of two highly
distinctive types, one of araucarian and the other probably of taxodiaceous affinity :
and in the case of the latter the foliation of the cone pedicel may be superficially
compared with that of the separate leafy shoots, which have not been found in connec-
tion with cones. The foliated and defoliated shoots may even include more than
one species amongst them, though at present these cannot be differentiated. The
name Araucarites sanctaecrucis is instituted for these sterile shoots, after the province
in which the petrified forest occurs.

(2) WOODS

(PI. i, figs. 2, 3, 7.)

These are represented by a large number of unbranched woody stems varying from
about 8 cm. to 0-5 cm. in diameter, a few being still partially embedded in the rhyoli-
tic ash (PI. i, fig. i, w). Most of these specimens have a complete thick cylinder
of secondary xylem, but no distinctive external features : PI. i, fig. 2 shows a stem
where part of an outer zone, probably representing remains of bark, may be distin-
guished (c). The preservation of the tissues, so far as examined, is so poor that
extensive cutting does not appear to be warranted : but it has been possible to make
a few observations on the wood structure. In the stem shown in transverse section
in PI. i, fig. 2, there is a narrow pith (p), and endarch primary xylem groups surroun-
ded by a wide zone of secondary xylem (x) with several somewhat indeterminate
growth rings, probably 12 in number. In radial longitudinal section the pitting of
the radial walls of the tracheids of the secondary xlyem can be observed only in a
few isolated patches where the organic matter of the walls has probably not been
entirely replaced during silicification : this is illustrated in PL i, figs. 3, 7. The
bordered pits are uniseriate and contiguous with slight flattening of outline where
they adjoin ; but the structure of the medullary rays has nowhere been observed.
In another larger specimen (8-5 cm. in diameter : V 3°976), probably from a branch
16 years old as judged by the ill-defined growth rings, similar uniseriate pitting has
been observed, but again without preservation of medullary ray structure. This
wood type so far as it can be defined is not identical with that reported by Gothan

106 A CONIFEROUS PETRIFIED FOREST IN PATAGONIA

(1925 : 198, pi. i, figs, i, 2) and Wieland (1935 : 16) in other specimens from the
same locality. Gothan's wood, named by him Dadoxylon (Araucarioxylon) sp., show-
ed radial tracheid walls with uniseriate bordered pits, sometimes distantly separate
and rounded, sometimes contiguous, but with the indication of the cross-field pitting
too indefinite for satisfactory use.

Uniseriate contiguous pitting of the tracheids such as is illustrated here was a
type widely represented in the Mesozoic (cf. Seward, 1919 : 165 ; Krausel, 1949)
by a variety of " genera " of fossil woods e.g., Mesembrioxylon and Brachyoxylon.
It has from time to time been interpreted as one transitional phyletically between
the " pinacean " and " araucarian " types as these were formerly more or less
rigidly defined (e.g., Hollick & Jeffrey, 1909 : 75) : Wieland regarded the Cerro
Cuadrado woods as representing such a transitional type (1935 : 16). However,
it has more recently been demonstrated that this type of xylem pitting cannot be
regarded as necessarily indicative of transitional or ancestral Araucariaceae, since
it is found within the range of variability in xylem structure in individual plants
in modern genera of different families of the Coniferales (e.g., Pinus : Bailey, 1933 ;
Sequoia : Bailey & Faull, 1934 ; Araucariaceae : Pool, 1929). According to Pool
(1929 : 599) it is more common in the stem wood of Araucariaceae than is usually
recognized.

Without more details of the wood structure, therefore, and in particular of the
medullary rays on which keys for wood identification have laid stress (Gothan, 1905 ;
Krausel, 1919, 1945 ; Phillips, 1941), it is not possible to make any effective com-
parison or identification of these woods. As will presently be shown, they are asso-
ciated with two types of cone, one araucarian and the other probably most nearly
related to the Taxodiaceae ; and the wood just described might have belonged to
either-it corresponds closely with that described for the cone axis of the latter type.
Similar wood structure has also been found, very poorly preserved, in the sterile
foliated branches of araucarian habit already described. But it cannot be identified
more specifically than as coniferalean wood.

It may be noted in conclusion that Wehrfeld (1935) referred to abundant remains
of araucarian petrified wood in the forest, but he adduced no evidence of structural
detail. His illustrations are of uncut woody trunks and branches, some showing
" annual " rings (e.g., p. 120, pi. f), and some of the trunks were very large, one
incomplete specimen being reported as 100 metres in length (p. 125). If this were
confirmed, it would seem that some of the trees of the Cerro Cuadrado petrified forest
were perhaps the tallest that ever lived, and certainly the tallest that have been
found in a petrified state. One would like to have precise measurements, not only
of length, but of diameter at intervals throughout the length. As Fossa-Mancini
has pointed out (1941 : 75), the theoretical maximum height of about 300 feet
propounded by Galileo for a tree (cf . Thompson, 1942 : 28) was no doubt considerably
exceeded by such a tree as that mentioned by Wehrfeld : and other reports (Feruglio,
1949 : 129) have also recorded very wide trunk bases in the forest. Leanza is
quoted by Feruglio (1949 : 129, footnote) as reporting the base of a fallen trunk of
circumference 10-3 metres (i.e., diameter approx. 3-26 metres). These accounts
of gigantism in the trees of the petrified forest refer to the trunks as araucarian ;

A CONIFEROUS PETRIFIED FOREST IN PATAGONIA 107

as there is no account of wood structure to corroborate this, the identification has
no doubt been made on the evidence of associated cones. The wood structure of
branches so far examined structurally, as has been pointed out, is indefinitive in
character for purposes of identification ; and as will later be shown, while one of the
cone types found in the forest is araucarian, the other is probably most nearly related
to the Taxodiaceae, in which family at the present day occur the most striking exam-
ples of gigantism in trees, comparable with those of the petrified forest. It is perhaps
more likely that these giant petrified trunks belong to this family than to the Arau-
cariaceae, but more information on the details of their wood anatomy is needed.
Wehrfeld also reported finding, more rarely, wood of palms. His description of
these mentions woods showing yellowish spots which he interpreted as the remains
of the principal vascular bundles ; but no structural evidence was given to support
this interpretation, nor has it so far been possible to obtain evidence of the presence
of monocotyledons from any other woods examined. The " yellowish spots " which
he described may represent variation of the mineral differentiation in the stems rather
than the position of vascular bundles ; for the petrifactions in the forest tend to
show differentiation of colours in the quartz.

(3) SEEDLINGS

(PI. 2, figs. 14-25 ; Text-fig, i, A-C.)

There are numerous (about a couple of hundred) petrified structures, many of them
looking like small corms or carrots, which vary in shape from ellipsoidal to turbinate
(PI. 2, figs. 14, 15, 17,), with intermediate forms such as that shown in PI. 2, fig. 16.
Most of these specimens are detached : but PI. 2, fig. 14 shows one (s) lying prone
and partially embedded in the matrix. The turbinate forms vary in size from 4 cm.
in height and 5 cm. in widest diameter to 1-5 cm. in height and 2-2 cm. in diameter ;
the ellipsoidal or clavate forms are more uniform in size, averaging 4-5 cm. in height
and 1-5 cm. in widest diameter. None of these structures shows any continuation
at either end of the axis into a leafy shoot or a well-defined root : PI. 2, figs. 18 and
19 show one specimen with a prolongation of the broader end of the axis which
bears no evidence of insertion of lateral organs.

From external examination of the specimens and from examination of cut and
polished surfaces it may be seen that most are partially or completely decorticated
(PI. 2, figs, 15, 17), and that a few have what appears to be a fairly thick periderm
(PI. 2, figs. 18-20, 22, pd.}. The core of such specimens is composed of a thick
zone of poorly preserved tissue resembling secondary wood (PI. 2, figs. 19, 20, 22, x)
and a narrow region in the centre probably comprising pith and primary xylem.
The decorticated specimens show on their external surfaces longitudinal fissures and
markings (PL 2, figs. 15-17), some of which might indicate the position of primary
medullary rays in the xylem ; however, as such markings may also be found on
corticated specimens (PI. 2, fig. 18), they may be due, at least in part, to cracking
during decay and preservation.

Some thin sections have been prepared from the more promising of these specimens
with a view to studying the arrangement and detailed structure of the tissues for

io8

A CONIFEROUS PETRIFIED FOREST IN PATAGONIA

comparison with the gross morphology. The preservation is unfortunately very
poor in most of the specimens cut : but PI. 2, figs. 23-25 shows the centres of trans-
verse sections taken at the top (broader) end, middle, and base (narrower end) of a
specimen similar to that shown in PI. 2, fig. 15, where preservation was better than
in most. The outlines of the whole sections are shown in Text-fig. I, A-c (correspon-
ding with PI. 2, figs. 23-25), where it can be seen that there is a medullated stele
with the pith region (p) widening from the base upwards, whilst it also changes from
rounded to ovate in shape in transverse section. PI. 2, fig. 21 shows this pith in
median longitudinal section of the broader end of the specimen ; the innermost

TEXT-FIG, i. Seedling (cf. Araucaria mirabilis). Transverse sections through a seed-
ling at top, middle and base of swelling (cf. PI. 2, fig. 15), showing pith (p) in centre
surrounded by primary xylem groups (pr) and secondary xylem (sec.), (cf. PI. 2, figs.
23-25)- V. 30951. x 5.

elements of the xylem ring are narrower than those forming the bulk of the xylem,
and the pith is notable for its more or less isodiametric, thick-walled cells. The
bulk of this specimen is composed of poorly preserved tissue resembling secondary
xylem traversed by medullary rays, in which structure of pits cannot be distinguished.
The primary xylem groups which abut on the pith increase in number from below
upwards, and the condition appears to be endarch, with stem structure throughout,
and with no evidence of traces passing out to lateral appendages. Other sections
from specimens similar to those in PI. 2, figs. 16, 17 show the same general arrange-
ment of the tissues (cf. PI. 2, fig. 22) ; the structure of the tissue resembling secondary
xylem can only be distinguished in a few better-preserved patches in some of the

A CONIFEROUS PETRIFIED FOREST IN PATAGONIA 109

longitudinal sections, where coniferalean pitting, of the type illustrated in PI. i,
figs. 3 and 7, may be distinguished on the radial walls.

These curious bodies must therefore be interpreted as swollen intercalary organs,
having stem structure but without any evidence of lateral appendages, and with
marked development of a tissue resembling secondary xylem. Their identification
is a matter of some difficulty ; but the structures are so numerous, and the inter-
calary swelling is so striking in all of them, that they must have some regular signi-
ficance in the vegetation of the forest. Wehrfeld figured a number of such structures
(1935 : 129), and referred to them as " a scientific novelty — a series of fossil arau-
carias," without, however, substantiating in any way the reference to the Araucaria-
ceae. Gothan, too, in a recent brief study (1950), has described such structures as
those in PI. 2, fig. 17 — the turbinate or fig-shaped forms — and believes the most
likely interpretation to be that they are of the nature of insect galls, terminal on
branches, where hypertrophy of the wood of the branch has been caused with the
larva living in the centre, and with the gall eventually dropping off the end of the
branch. He also believed they might be araucarian, but had found no evidence of
any such galls in living Araucariaceae. It has not, however, been possible to find
any evidence that the structures are other than intercalary, or that the turbinate
forms are different in kind from the ellipsoidal forms ; and in some of the specimens
sectioned there is a definite pith including thick-walled cells and with no sign of
any larval intrusion. There remains, however, the possibility that these might
be fungal stem galls of an intercalary nature ; such intercalary stem swellings are
known in some cases to be caused by fungi, as in the case of Peridermium galls on
Pinus stems ; but no records of such galls on araucarian stems have been found,
though Connold (1909 : 132, text-fig. 159) has described long, irregular tumours on
the roots of Araucaria imbricata which do not resemble the present structures at
all. In the case of either terminal or intercalary stem galls, however, one would
expect to find some evidence of lateral appendages, and there is none. On the other
hand the size, shape and internal structure of these bodies all tend to emphasize
their similarity to broken first-year seedlings of the genus Araucaria in the sections
Bunya and Columbea, where tuberous development in varying degree is found in the
hypocotyl, which is of stem structure with early incidence of secondary thickening
(Diirr, 1865 : 103 ; Hemsley, 1901 ; Seward & Ford, 1906 : 333 ; Shaw 1909 ; Hill
& de Fraine, 1909 : 212 ; Hickel, 1911 : 160).

Wieland has figured certain branch-like structures (1935 : 27, pi. 13, fig. 2) as
bases of second-year seedling stems of Proaraucaria, with broken root bases : but
if these are indeed older seedlings, they are not markedly tuberous, and there is
no evidence of any relationship with the smaller structures here described. The
tuberous habit in seedlings in general is sporadic : it appears to be rare amongst
gymnosperms, occurring in the Cycadales in Encephalartos , and in the Coniferales
in the two sections of the genus Araucaria just mentioned. The proximity of these
numerous tuberous bodies to fertile cones belonging to the section Bunya of the
genus Araucaria, which will be described in the next section, also tends to corroborate
their interpretation as araucarian seedlings : and the fact that the embryos in the
seeds of these cones are dicotyledonous might suggest that the markedly ovate form

no A CONIFEROUS PETRIFIED FOREST IN PATAGONIA

of the pith at the upper end of the seedlings is related to a stelar adjustment below
the level of the insertion of the vascular supply of two cotyledons on opposite sides
of the hypocotyl axis.

So far as has been ascertained, this is the only record of petrified seedlings. Kendall
(1949, text-fig, i, j.) has drawn attention to a curious specimen, found as a compres-
sion, from the Mid- Jurassic Estuarine Series of Yorkshire which she has interpreted
as a seedling stage of Amucarites phillipsi Carruthers, but no other such records are
known to the writer.

Mr. W. N. Edwards informs me that there is a specimen in the Sedgwick Museum,
Cambridge, from the Lower Greensand of Upware, Cambridgeshire, which was
figured by Keeping (1883 : 150, pi. 8, fig. 7) as " a small fruit (?) ferruginized,"
but which superficially at least is very similar to some of the smaller araucarian
seedlings from Patagonia.

(4) CONES

The cones, which represent the best preserved part of the collection, are of especial
interest, since petrifactions of reproductive organs of fossil conifers are seldom found,
though petrified coniferous woods are abundant. Two distinct types of seed cone
are represented, but no male cones or pollen-producing organs have been found.
These two types of cone have already been partially described by Spegazzini (1924),
Gothan (1925), and Wieland (1935) ; but detailed studies of the structure, which
affect the interpretation of the morphology and consequent assessment of the relation-
ships of the cones, have not been made, with the exception of Darrow's investigation
(1936) of the structure of the embryo of Araucaria mirabilis.

Araucaria mirabilis (Spegazzini)
(Pis. 3, 4, 5 ; Text-figs. 2-4)

1924. Araucarites mirabilis Spegazzini, p. 126, text-figs. 1-3, 4 (1-7).

1925. Araucaria windhauseni Gothan, p. 200, pis. 2-7, pi. 8, fig. I.
1929. Proaraucaria mirabilis (Speg.) Wieland, p. 60.

1929. Proaraucaria elongata Wieland, p. 60.

1931. Araucaria mirabilis (Speg.) Windhausen, p. 201.

1935. Proaraucaria mirabilis (Speg.) : Wieland, p. 19, pi. i ; pi. 7, figs. 2, 3 ; pi. x ; pi. xi, figs.

i, 2, 4 ; text-fig. 4.

1935. Proaraucaria mirabilis var. elongata Wieland, p. 26, pi. 8, fig. i ; pi. 9, fig. 2 ; pi. 12, fig. 4.

1935. Proaraucaria patagonica Wieland, p. 26, pi. 6, figs. B, D, E, F.

1935. Proaraucaria mirabilis var. minima Wieland, p. 26, pi. 6, fig. A ; pi. 12, fig. 3.

1936. Proaraucaria mirabilis (Speg.) : Darrow, p. 328, text-figs. 1-13.
1936. Araucarites mirabilis Speg. : Gordon, p. 14.

1939. Proaraucaria mirabilis (Speg.) : Darrah, p. 222, text-figs. 133, 134.

1940. Araucaria mirabilis (Speg.) : Florin, p. 36.
1944. Araucaria mirabilis (Speg.) : Florin, p. 513.

1947. Proaraucaria mirabilis (Speg.) : Arnold, p. 313, text-fig. 156.

1947. Proaraucaria mirabilis (Speg.) : Andrews, p. 177, text-fig. 121.

1948. Proaraucaria mirabilis (Speg.) : Wilde & Eames, p. 312.

1949. Proaraucaria mirabilis (Speg.) : Feruglio, p. 129.
1951. Proaraucaria mirabilis (Speg.) : Feruglio, p. 35.

A CONIFEROUS PETRIFIED FOREST IN PATAGONIA in

EMENDED DIAGNOSIS. Seed cones varying in shape from spherical to somewhat
ellipsoid, and in size from about 8 to 4 cm. in height and 8 to 4 cm. in diameter.
Cone-scales numerous, arranged in a close spiral, one-seeded, with thick woody wing
and deciduous laminar tip, which on dropping reveals the large woody ligule on top
of the rhomboidal apophysis of the bract ; the fertile scale is fused with the bract
for about two-thirds of its length, so that the depth of the ligular sulcus is about
one-third of the length of the fertile scale. Cone-scales of mature seed cones (de-
laminated) 13-16 mm. long and 10 mm. wide including the wing, tapering somewhat
towards the base. Ligule 4 mm. wide, 1-2 mm. high, about 5 mm. long. Seed
8 mm. long and 3 mm. wide and deep, inverted, albuminous, apparently embedded
in the fertile scale tissue ; testa with a thick stony layer. Embryo large, dicotyle-
donous, with numerous spirally twisted suspensors at base. Evidence on mode of
seed dispersal lacking.

Axis of cone wide, with a wide pith including branched sclereids, surrounded by a
narrow ring of separate vascular bundles each with a wide outer sheath of fibrous
extra-phloem tissue. Vascular supply of cone-scale double, the bract trace single
and arising separately from the base of the leaf-gap in the axis stele, the fertile scale
supply of two strands, arising from either side of leaf gap, which fuse with inversion :
both bract and fertile scale supply fork in base of cone-scale to give a lower and an
upper (inverted) series respectively, the lower series passing out to the apophysis and
laminar tip of the bract accompanied by a series of resin canals, the upper series
terminating in the large strongly vascularized ligule.

DESCRIPTION. — The species is represented by a large number of specimens in the
collection ; most of these are isolated specimens, but a few are still partially em-
bedded in the matrix (PI. 3, fig. 26). Some of the cones must have been mature,
as they contain fully formed embryos in the seeds, while others have no embryos,
though the cones may be as large as those which do. The cones vary in size from
8 cm. in length and diameter to 4-4 cm. and 4 cm. in length and diameter respectively,
and in shape from nearly spherical to ovoid or ellipsoid. There is no evidence to
suggest that the smaller cones are necessarily immature, for one about 5 cm. in
length and diameter (V. 30975) has fully formed embryos in the seeds. Some
specimens appear more markedly elongate, with an ellipsoidal section, than others
(e.g., PI. 4, figs. 38, 42) : such cones were interpreted by Wieland (1935 : 26) as a
distinct variety, Proaraucaria mirabilis var. elongata, but structurally it has not
been possible to differentiate these types, and there is evidence from the cone figured
here that the condition has been exaggerated by considerable crushing prior to petri-
faction, as may be seen in the compressed and broken outer parts of the scales in
PI. 4, figs. 40, 42, while the proximal part of the scales has withstood the compression,
being reinforced, perhaps, by the woody tissues of the testa. The cones have been
petrified before the shedding of the seeds in most cases ; no separate petrified seeds
nor cone-scales have been found, but there are a few specimens, such as those figured
by Wieland (1935, pi. 6, E) and interpreted by him as another species, Proaraucaria
patagonica, which no doubt represent the naked axes of the cones after the cone-
scales have been shed (e.g., V. 30969). They correspond in size with the measure-
ments of the cone axes themselves ; one has been sectioned, and shows a wide pith

112 A CONIFEROUS PETRIFIED FOREST IN PATAGONIA

with sclereids, and peripheral vascular bundles. The surface features correspond
closely with those found in cones of living araucarians after shedding of the cone-
scales (cf. Seward & Ford, 1906 : 361), with a reticulum of low ridges outlining rhom-
boidal areas no doubt representing the areas of attachment of the scale bases to the
cone axis. The absence of any separate cone-scales, which Gothan (1950 : 153)
has emphasized as a feature suggesting that these cones did not shed their scales
at maturity, and as one which might separate them genetically from those of other
species of Araucaria, may well be explained in terms of the season in which the
forest was overwhelmed. The seeds in the cone-scales in the final year of growth
had not been shed, and the previous year's had germinated and are probably repre-
sented in the forest by the very numerous petrified seedlings to which reference has
already been made.

Various views of external features of cones are shown in PI. 3, figs. 26-31, PI.
4, figs. 38, 39, and PL 5, fig. 46. Inserted on the cone axes are a large number of
close-set ligulate cone-scales, each of which bears a single large inverted seed sunken
in its tissue on the upper surface. The distal, abaxial faces of the cone-scales are
seen arranged in a close spiral forming some 55 orthostichies (so far as has been
examined). The surface morphology of the cone-scales varies, probably according
to the age and condition of the cone and also to some extent to the degree of weather-
ing of the surface, as shown in PI. 3, figs. 28, 29, 31, 34, PI. 4, figs. 38, 39, and PI. 5,
fig. 46. PI. 4, figs. 38, 39, 42 show a cone where in surface view the cone-scales
have laminar, longitudinally striated tips (t). In earlier descriptions of the cones,
Gothan (1925, pi. 2) and Spegazzini (1924, text-figs, ib, 4 (6)) have illustrated some
with a similar triangular lamina on the cone-scales, and Wieland mentions this
feature (1935 : 175, pi. 10, fig. 2). Most of the cones, however, show surface mor-
phology of cone-scales as illustrated in PI. 3, figs. 28, 29, and PI. 5, fig. 46, with
prominent apophysis of bract (a) with marked lateral wings (w), and a large ligule
(/). Consideration of sections of the specimen in PI. 4, figs. 38, 42, leaves little doubt
that this latter condition is the result of the dropping of a laminar tip, though erosion
of surface features may occasionally account for it. PI. 4, figs. 40, 42, show this
laminar tip t, covering the ligule /, in radial longitudinal section ; and Text-fig. 4,
A-I, and PL 4, fig. 43, PI. 5, fig. 54, from a tangential longitudinal series, demon-
strate the same relationship between the laminar tip and the ligule. There is, too,
some structural evidence of an absciss zone at the base of the laminar tip, e.g., at
A in PI. 4, fig. 40. The lateral wings of the cone-scales are also seen to be present
in the series of tangential sections, though they tend to be obscured in surface view
(PI. 4, fig. 39) as compared with the cones with delaminated scales. In any respect
other than the presence or absence of a laminar tip it is difficult to separate the two
cone types : in both, seeds with fully formed dicotyledonous embryos of the same
type have been found. The much smaller cone illustrated in PI. 3, figs. 31-34, shows
similar longitudinally striated laminae clothing the exterior of the cone ; in section
(PI. 3, figs. 32, 33) no ovules can be distinguished, and it may be either a young
cone or one with abortive or unpollinated ovules. The comparatively large ligule
can again be distinguished behind the distal lamina of the cone-scale (PI. 3, fig. 33,/).
There seems little doubt that the three cone types just described represent

A CONIFEROUS PETRIFIED FOREST IN PATAGONIA 113

different stages in the development of the same cone species ; in many of the living
Araucariaceae there is a laminar tip or spine on the cone-scale which often drops off
when the cone is ripe (cf. Pilger, 1926 : 255).

The ligulate cone-scales are modified in form towards the base of the cone, where
they are sterile. PI. 3, fig. 30 shows the transversely cut tip of a cone pedicel (pi),
around which are a number of such sterile cone-scales of simplified form, which
pass over gradually into the fully formed fertile cone-scales above. This is a charac-
ter of araucarian cones which has been emphasized by Seward & Ford (1906 : 354,
text-fig. 22A) as contrasting with the condition in the Pinaceae where the cone-scales

b.s.

^m^i \

B

b.s. —

TEXT-FIG. 2. Araucaria mirabilis (Spegazzini). Transverse sections through vascular
bundles of cone axis stele, showing from base upwards a series in the detachment
of the bract supply (b.s.) and fertile scale supply (/i, /2) of cone scales. The two traces
of the fertile scale supply fuse with inversion to give a single bundle (f.s.). The pith
lies to upper side, cortex to lower side, of the vascular bundles, where xylem is shown
black, with protoxylem white, phloem finely stippled, and extra-phloem tissue
coarsely stippled. V. 30958. x 27-5.

are much more sharply differentiated in form from the foliage leaves, and succeed
them more abruptly. In none of the fossil cones examined is there critical evidence
of the foliation of the pedicel, or of attachment to any of the leafy branches which
are found amongst the remains of the forest.

The axes of the cones are thick (PI. 3, fig. 27), with a wide pith and narrow ring
of separate vascular bundles, seen at p and v respectively on the transversely frac-

GEOL. II, 2. 8

114 A CONIFEROUS PETRIFIED FOREST IN PATAGONIA

tured surface and in Text-fig. 2 in transverse section. The pith contains branched
sclereids ; the tissues of the vascular bundles are ill-preserved, but in patches of
better preserved tracheids the xylem pitting varies from uniseriate to biseriate and
alternating, with the pits separate, and each bundle is endarch, with a narrow zone
of phloem, outside which is a deep arc of extra-phloem tissue (? pericycle) as seen in
Text-fig. 2, e.p. This extra-phloem tissue is much better preserved than is the
phloem, and consists of large, thick-walled, sometimes branching cells resembling
the sclereids of the pith rather than a typical fibrous tissue. Comparison of this
type of cone axis with that of other conifers reveals the striking similarity with that
of living araucarian cones. Thomson (1913 : 4) has drawn particular attention to
the exceedingly large size of the pith in the cone axis of the Araucariaceae, especially

TEXT-FIG. 3. Araucaria mirabilis (Spegazzini) (cf. cone in PI. 4). Tangential longitudinal
sections of cone-scales from base outwards, showing splitting of the two bundles b.s.,
f.s., in base (cf. Text-fig. 2), to give the vascular supply of bract and fertile scale respec-
tively. Xylem shown black, with protoxylem white, extra-phloem tissue coarsely
stippled. V. 30957. x 27-5.

in the seed cones, where it may reach i in. in diameter. The sclereids of the pith
and the extra-phloem or fibrous pericyclic tissue are, too, characters associated with
the axis anatomy of the living Araucariaceae (cf. Seward & Ford, 1906 : 337).

A study of the detachment of the cone-scale vascular supply from the axis stele
has been made in transverse sections of the cone axis ; Text-fig. 2, A-F, illustrates
successive stages, from below upwards, in the detachment of the traces, as seen in
different sectors of .t he wide cylindrical stele. This has proved more satisfactory than
the use of a series of sections, as 1-5-2 mm. may be lost between successive sections
in preparing a series, while the spiral succession of the numerous scales is so close-set
that from one section a close series of stages can be obtained. The bract supply ;

A CONIFEROUS PETRIFIED FOREST IN PATAGONIA 115

b.s., comes off as a single trace from the base of the leaf-gap, and free from the fertile
scale supply, f.s., which is derived from the fusion of two strands, /i, /2, detached
separately from either side of the leaf-gap above the bract supply. These two
strands rotate through 180° while fusing, so that the resultant fertile scale fusion
bundle is inverted (Text-fig. 2, D-F). In the base of the cone-scale there are, there-
fore, two bundles, the upper one inverted. Text-fig. 3 shows their subsequent
behaviour in tangential sections of the cone illustrated in PI. 4, fig. 38, taken in
series from the inside of the cone outwards : Text-fig. 4, from the same series, shows

H

TEXT-FIG. 4. Araucaria mirabilis (Spegazzini) (cf. cone in PI. 4, and Text-fig. 3).
Tangential longitudinal sections of cone-scales from base outwards, to show vascu-
larization of bract (b), its laminar tip (t], and fertile scale (/) with ligule (/). Xylem
shown black, sclerotic tissue coarsely stippled. S — seed, T.S. = stony layer of testa,
e = embryo, d — resin ducts, f.s. = vascular supply of fertile scale, b.s. = vascular
supply of bract. V. 30957. x 5.

the relationship of the bundles to the cone-scale morphology throughout the scale
and into the laminar tip. Each of these two bundles in the scale base branches, the
bract bundle first (Text-fig. 3, D-E), and this lower series, though incompletely pre-
served in the region below the seed (Text-fig. 4, E, F), appears to supply the apophysis
and the laminar tip of the bract (Text-fig. 4, G, H, I ; cf. PI. 4, fig. 43 ; PI. 5, fig.

n6 A CONIFEROUS PETRIFIED FOREST IN PATAGONIA

54). In the outermost region of the cone-scale (Text-fig. 4, G, H, and PL 5, fig. 54)
structures resembling resin canals (d) lie below the bundles. PI. 4, figs. 43, 45 show
obliquely cut bundles (v.b.b.) supplying the laminar tip of the bract ; these bundles
are accompanied on either side by a strand of cells with dark contents (d), which
may represent the upward extension of the resin canals into the laminar tip. It
has not been possible to demonstrate whether, as in Araucaria bidwilli, there is a
distinct system of bundles ending blindly in the apophysis, with the laminar supply
branching off farther back.

The upper inverted fertile scale bundle branches farther out in the scale base
(Text-fig. 3, E-G), and forms a strong inverted upper series, probably supplying the
base of the single inverted seed (Text-fig. 4, F) as well as the large ligule (Text-fig. 4,
G, H, and PL 5, fig. 54). PL 4, fig. 44 demonstrates a vascular strand in the ligule.
PL 5, fig. 53, taken from a delaminated cone, also demonstrates vascularization of the
large ligule (/), along with the distinct series of the bract (b).

There are two notable features here for comparison with the living species of
Araucaria : the separate origin of the bract and fertile scale supplies, and the strong
vascularization of the ligule. Only in the section Bunya Wilde & Eames amongst
the living araucarians are these two conditions found (Eames, 1913 : 24 ; Aase,
1915 : 297 ; Wilde & Eames, 1948 : 322 ; 1952 : 44). This is probably the first time
that it has been possible, in a fossil araucarian cone, to demonstrate these characters,
which are of critical importance in considering the taxonomy and interrelationships
of living araucarians.

Text-fig. 4 and PL 5, fig. 54, also serve to demonstrate the lateral winging of the
cone-scale ; this " winging," already noted in the external morphology of the cones,
extends from near its base out to the apophysis, as seen in Text-fig. 4, B-H. A similar
condition is found in the delaminated cone shown in PL 5, figs. 46, 50, 53. The wing
is about as wide as the body of the seed, and is relatively thick ; it is supplied by
vascular bundles of the lower (bract) series (Text-fig. 4, G, H, and PL 5, fig. 54). The
tissues of the wing have evidently been woody, as may be seen in the cone in PL 5,
fig. 50, where a core of thick-walled pitted cells is found, possibly of the nature of a
transfusion tissue, and in PL 5, figs. 53, 54 where there is a deep hypodermal zone of
thick-walled cells. Cone-scales as thick and woody and as widely winged as these
must have been are found amongst living araucarians only in the section Bunya
(Wilde & Eames, 1952 : 44).

The morphology of the ligule, already noted in surface view of the cones, is seen
in tangential longitudinal sections in Text-fig. 4, G, H, PL 5, figs. 53, 54, and in radial
longitudinal section in PL 4, fig. 40, PL 5, fig. 52. Its vascular bundles have already
been noted ; its ground tissue is similar to that of the wings, and no doubt it was also
woody. The depth of the sulcus between the ligule and the bract may be exaggerated
by artificial splitting between the two, extending apparently deep into the scale,
the space occupied by clear quartz. The cone in which the split is most clearly and
perhaps most naturally seen is illustrated in PL 3, figs. 35-37 : it will be noted that,
where the scale is not quite medianly cut as at A in the radial longitudinal section
of the cone in PL 3, figs. 35, 36, the split extends deeper than the level of the base
of the seed, but where a more median cut of the seed is found, as at B, the split stops

A CONIFEROUS PETRIFIED FOREST IN PATAGONIA 117

towards the base of the seed. This is also demonstrated in the tangential longitudi-
nal series of another cone in Text-fig. 3, F-H, where in Text-fig. 3, G, just outside the
base of the seed, the ligule has separated from the bract on the margin but not yet
in the median plane. It will be seen from PI. 3, figs. 35, 42 that the ligular sulcus
is about one-third the depth of the fertile scale, the fertile scale being fused with the
bract for about two-thirds of its length ; and this is about the same proportion for
the depth of the ligular sulcus as in the living Araucaria bidwilli in mature cones
(cf. Wilde & Eames, 1948 : 325 ; Pilger, 1926 : 256, text-fig. 139, A). Wieland
based his generic definition of these cones as Proaraucaria on the greater depth and
size of the split separating ligule from bract, as well as on the larger size of the ligule,
in comparison with the condition in living species of the genus Araucaria (Wieland,
1935 : 19), and he regarded the fossil species as representing an intermediate stage
in the evolution of the cone-scale, with less complete fusion of bract and fertile
scale. His figures, however, nowhere make clear the natural depth of the split
separating ligule from bract. From the present examination there is no evidence
to show that the difference in degree of size of ligule and depth of sulcus separating it
from the bract in the fossil and in living species is such as to warrant the erection
of a new genus ; the fossil is closely similar to Araucaria bidwilli in these respects.

The relationship of the seed to the upper surface of the scale is shown in Text-
fig. 4, B-G, PL 4, figs. 40-42, and PL 5, figs 47, 50-52. The single large inverted
seed (s) appears as if embedded in the scale tissue ; this is probably best seen in the
tangential sections in PL 5, figs. 47, 50, the conspicuous stony layer of the testa
(T.S.) lying within an outer sheet of tissue (F) which is continuous with the surface
tissue of the wing of the scale on either side of the seed. Within the stony layer the
nucellus is imperfectly preserved, but in many seeds the embryo sac may be defined,
enclosing the endosperm with an embryo embedded in it (PL 5, figs. 47, 50-52, en
and E). The dicotyledonous embryo is seen entire in PL 4, fig. 41, where the polished
radially cut surface of the cone is photographed ; and embryos are seen cut in trans-
verse section of the cone at E in PL 5, figs. 48, 51, and Text-fig. 4, E, in tangential
longitudinal sections of the cone in PL 5, figs. 47, 49, 50, and in radial longitudinal
sections in PL 4, figs. 40-42 and PL 5, fig. 52. Plate 4, fig. 41 and PL 5, figs 48 49
demonstrate the two cotyledons (co), and PL 4, figs. 41, 42, show the orientation
of the embryo with the thick radicle pointing towards the micropylar end of the em-
bryo sac, where there is a tangled mass of suspensors (ss) (PL 5, fig. 51). The embryo
need not be described in detail as this has been done by Darrow (1936) ; but dico-
tyledonous embryos of similar type have been found in both the laminated and
delaminated types of cone (cf. PL 4, fig. 42, PL 5, figs. 47-51), and the orientation
of the plane in which the median lines of the two cotyledons lie varies from one more
or less parallel to the surface of the fertile scale (PL 5, figs. 47, 49) to one at right
angles to it (Text-fig. 4, E).

RELATIONSHIPS. From the fuller account of these araucarian cones which it
has now been possible to give and with particular reference to the critical characters
of mode of origin of the vascular supply of the cone-scale from the axial stele, size
and nature of wing of cone-scale and size and nature of ligule, two major conclu-
sions emerge which contrast with many of the previously expressed views on the

n8 A CONIFEROUS PETRIFIED FOREST IN PATAGONIA

relationships of this species. The first is that there are no grounds for its taxonomic
separation into a new genus Proaraucaria, which was effected by Wieland (1935)
because he believed the cleavage between ligule and bract to be deeper than in any
living species of Araucaria. Gothan later (1950 : 153) suggested, without further
description of the cones, that the fact that they did not appear to shed their cone-
scales at maturity was a character of major importance in separating the genus
Proaraucaria : this condition has been discussed in an earlier section, where it was
concluded that there is no evidence that the cones did not shed their scales, but a
certain amount to suggest that they did. The second major conclusion is that the
nearest related living species of Araucaria is A. bidwilli, the only species so far inclu-
ded in the section Bunya recently erected by Wilde & Eames (1952) in their taxo-
nomic revision of the genus Araucaria, and found to-day only in Eastern Australia
in Queensland. Most of the previous references to these fossil araucarians have
compared them with the two living South American species, A. araucana and A.
angustifolia ; though Darrow compared the fossil cones with A . bidwilli, this was
not substantiated by any critical study of the vascularization or of the ligule, but
she effectively demonstrated the structure of the embryo, which is similar to that
in A. bidwilli. It is true that in the fossil species there is no evidence for dehiscence
of the individual cone-scales, for no separate seeds nor empty scales have been found ;
and this character is one of those on which Wilde & Eames differentiated their new
section Bunya (1952 ; 44). But in general structure of the cone-scale and ligule,
and in particular of their vascularization, there is a striking similarity between the
living Bunya and the fossil species : indeed, the larger size of the cones of A . bid-
willi, which may reach 30 cm. in diameter, appears to be the chief difference between
the species as represented by the cones. The use of the generic name Araucarites
for these cones, though it has taxonomic priority, is best set aside (cf. also Florin,
1940 ; 1944) : for this name is reserved for cones or branches of araucarian habit
which, however, are lacking in preservation of structural details which might give
final evidence justifying inclusion in the genus Araucaria itself. The name Arau-
caria mirabilis (Speg.) is therefore used, and the species is assigned to the section
Bunya Wilde & Eames, the only other species of this section being the living
Araucaria bidwilli, which is found to-day only in Queensland.

The proximity of these cones to the seedling structures already described is certain-
ly significant, though there is no evidence of organic relationship. Tuberous seed-
lings are known amongst living species of Araucaria only in the section Bunya and
in the South American section Columbea. The rarity of such a seedling habit in
gymnosperms has already been noted ; and so far as records have been examined
it is not found in any of the Taxodiaceae, to which the only other fertile cones in this
area where the seedlings are found probably belong.

Similarly, the proximity of the cones to the leafy branches of araucarian habit
which have been described as Araucarites sanctaecrucis may be of significance,
though evidence of attachment of cones to such branches is missing ; but as will
presently be shown, the foliation of the pedicel in the Pararaucaria patagonica seed
cones does not rule out the possibility that the branches, or some of them, may have
been related to this quite distinct type of cone.

A CONIFEROUS PETRIFIED FOREST IN PATAGONIA 119

Pararaucaria patagonica Wieland
(Pis. 6, 7 : Text-figs. 5-7)

1929. Pararaucaria patagonica Wieland, p. 60.

1929. Pararaucaria elongata Wieland, p. 60.

J935- Pararaucaria patagonica Wieland, p. 21, pis. 2-5.

1936. Pararaucaria patagonica Wieland

1937. Pararaucaria patagonica Wieland
1940. Pararaucaria patagonica Wieland
1944. Pararaucaria patagonica Wieland
1947. Pararaucaria patagonica Wieland
1949. Pararaucaria patagonica Wieland
1951. Pararaucaria patagonica Wieland

Gordon, p. 14.

Darrah, p. 223.

Florin, p. 36.

Florin, p. 513, pi. 184, figs. 23-26.

Arnold, p. 314.

Feruglio, p. 129.

Feruglio, p. 35.

EMENDED DIAGNOSIS. Seed cones, ovoid in shape, varying in length from 4-7
cm. to 2-3 cm. and in diameter from 2-4 cm. to 1-3 cm., borne on slender pedicels
clothed with spirally arranged, imbricate, broadly lanceolate and somewhat acute
leaves with longitudinal striation of the abaxial surface. Bracts bearing the axillary
fertile scales usually about 38 in number in average-sized cones, large and conspicuous,
probably woody, each subtending a thick fertile scale, and arranged in a closely
imbricate spiral succession with angle of divergence 3/8. Bract 10 mm. long, 12 mm.
wide, 1-5 mm. thick, free from fertile scale for greater part of length ; fertile scale
10 mm. long, 12 mm. wide, 2-5 mm. thick, showing longitudinal ridges on its pro-
truding distal abaxial face, and bearing usually one large inverted seed, laterally
inserted, flattened and winged, the seed separating from scale at maturity. Seed
6 mm. long, 6 mm. wide, 2 mm. thick, with wing 2 mm. wide tapering towards base
and tip of seed ; testa with inner stony layer, and outer layers composing wing
of a characteristic stellate sclerotic lacunar tissue. Embryo 4 mm. long, poly-
cotyledonary ; seed probably exalbuminous.

Cone axis slender, with narrow pith and thick endarch cylinder of xylem. Succes-
sion of elements in primary xylem includes scalariform and reticulate tracheids,
passing over into elements with biseriate bordered pits. Secondary xylem tracheids
usually with uniseriate bordered pits, contiguous and flattened, the pit apertures
rounded or ovate ; medullary rays with cross field pitting of cupressoid form. No
resin ducts or cells present. Bract supply arises from axis stele as a single trace
from base of leaf gap ; fertile scale supply arising as two traces, one from each side
of leaf gap above bract trace, the two traces fusing to give inverted bundle, which
is accompanied into the scale on the adaxial face by a large strand of sclerenchyma
which forks into two. Bract and fertile scale supplies single in base, branching
higher up, with an abundant transfusion tissue linking the bundles tangentially
and persisting in tips of bract and scale when bundles die out. Seed supplied by a
single lateral strand from supply of fertile scale.

DESCRIPTION. This species is represented by a large collection of mature seed
cones ; no younger stages appear to be present, though it has been possible to section
only a few of the specimens for more detailed examination. (One specimen, V. 3°974»
smaller than average with length 24 mm. and width 12 mm., probably represents an
abortive or unfertilized cone, but preservation is not good enough for a convincing

120 A CONIFEROUS PETRIFIED FOREST IN PATAGONIA

identification.) The species is redescribed here with special reference to a number
of diagnostic characters which were not included in Wieland's original account
(Wieland, 1935) ; it is the most interesting species in the collections from the Cerro
Alto localities, and its relationships with other conifers have not so far been assessed
by comparison of a sufficiently wide range of characters. It may be of interest to
note here that Feruglio (1949 : 129) reports, in a review of the localities where the
plants of the petrified forest are known to occur, that Pararaucaria. has been found
only in two of the localities with petrified cones, viz., the Cerro Alto, where it is
associated with the cones of Araucaria mirdbilis, and near the Estancia Los Toldos,
where the latter species is missing.

The general habit of the cones is illustrated in PI. 6, figs 55, 56, 63. They vary
in length from 4-7 cm. to 2-3 cm., and in diameter from 2-4 cm. to 1-3 cm., the average
size being about 4 cm. long and 2 cm. in diameter. There is no evidence that smaller
cones represent younger stages ; they probably represent attenuated cone forms,
bearing fewer appendages. This range in size corresponds with that observed by
Wieland in his specimens. A longer cone type was originally separated by him
(1929 : 60) as Pararaucaria elongata, but later (1935 : 21, pi. 5, fig. 6) he included this
type in Pararaucaria patagonica. A similar long cone is shown here in PI. 7, figs.
74, 75 ; and no other distinctive characters have been noted in it. These long
cones are often poorly preserved and much weathered, as in PI. 7, fig. 74, but the
bract and fertile scale show the same proportions as in smaller specimens. Cones of
average size as well as these long cones have been found to contain fully developed
embryos in the seeds (PI. 6, fig. 58 ; PI. 7, fig. 75), a condition refuting Wieland's
suggestion that the latter represent the mature seed cones, while the former are
immature. Such a range in size is a feature quite common in the cones of, for
example, living species of Pinus.

The cones are somewhat ovoid in shape ; many show marked unilateral weather-
ing of the surface features and some a varying degree of compression. The specimen
in PI. 6, fig. 63, approaches most nearly to the condition of the surface features in
the living plants. A number of large and conspicuous bracts (b), about thirty-
eight in an average-sized cone, are arranged in a close-set spiral around the axis,
with successive bracts overlapping and with an angle of divergence of 3/8 as in
other cones observed. In the axil of each bract is the fertile scale (/), showing
slight longitudinal ridging of the abaxial surface, where it protrudes distally.
Towards the cone apex the bracts and fertile scales diminish rapidly in size (PI. 6,
fig- 56), the topmost members being sterile (cf. series of sections V. 30965, V. 30968).
In some cones there is a markedly acute tip (PI. 6, fig. 56), in others it is more blunt
(PI. 6, fig. 55). Only two specimens among the cones have been found attached
to their pedicels : PI. 6, fig. 62, shows one of these, and clothing the pedicel is an
overlapping spiral series of broad, flat, lanceolate leaves, as seen in more detail in
PI. 6, fig. 64. These leaves are regularly striated on the abaxial surface ; the furrows
may represent the position of lines of stomata, but it has not been possible to verify
this from structural detail in either sections or cuticles. Another feature of some
diagnostic interest is also illustrated by this specimen : it shows a sharp transition
from the sterile leaves to the bracts and fertile scales at the base of the cone, a

A CONIFEROUS PETRIFIED FOREST IN PATAGONIA 121

character noted, for example, by Seward & Ford (1906 : 354) as one offering a
marked distinction between the Araucariaceae and the Pinaceae, the transition in
the former being much more gradual.

The narrow cone axis is seen in transverse section of the basal region of a cone in
PI. 7, fig. 65, and shows a relatively narrow pith and thick cylinder of xylem : the
xylem diminishes in thickness towards the tip of the cone. The pith includes large
scattered cells with dark brown contents : the stele is endarch, with scanty primary
xylem and a wide zone of secondary xylem with regular radial files of tracheids,
without parenchyma and without resin ducts or cells, and with unseriate secondary
medullary rays. The structure of the xylem is illustrated in more detail in PI. 7,
figs. 66-72. The centrifugal succession of elements in the primary xylem is shown
in radial longitudinal section from right to left in PL 7, figs 66-68. Annular tracheids
cannot be identified with certainty ; but a succession of spiral elements, in which
there is some evidence for the formation of bordered pits between the turns of the
spiral band (PI. 7, fig. 67, s.p.), is followed by widely reticulate tracheids (PI. 7,
fig. 67, ret.) and these are succeeded, in the base of the cone, by tracheids with bi-
seriate alternating bordered pits (PI. 7, figs. 70, 71, bi.), probably belonging to the
secondary xylem.

Beyond these come the tracheids forming the bulk of the woody cylinder ; these
have uniseriate bordered pits on the radial walls (PI. 7, figs. 68, 69, 70, sec.). The
pits are usually contiguous, with flattening of the pit outline in the area of contact
(fig. 69) : the form of the pit apertures varies from rounded to oval. Parts of the
medullary rays are seen in radial longitudinal section in PI. 7, figs. 68 — 70, 72 (m.r.)
The rays are 1-5 cells high and ray tracheids have not been observed. The cross-
field pitting is fairly constant (PI. 7, fig. 72), there being 2-4 horizontal rows of
rather small pits, which are of markedly " cupressoid " form (cf. Phillips, 1941:
267).

These characters of the xylem, not noted by Wieland, may most conveniently
be discussed here : they represent features which have been widely used in the past
as a means of identifying conifer woods.

Bailey has recently re-emphasized the comparative significance of the primary
developmental succession in the xylem of gymnosperms (Bailey, 1949, 1925 ; cf.
also Florin 1936, 1937) ; and in Pararaucaria the sequence which has just been
noted appears to be intermediate between that found in the " lower " gymnosperms
(Cycadales, Cordaitales) and that in the "higher" gymnosperms (Coniferales,
Ginkgoales, Gnetales). In the former groups scalariform elements are present and
are succeeded by reticulate elements, and these by elements with the typical bordered
pits of the metaxylem and secondary xylem ; bordered pits are not formed in the
spiral tracheids, where, however, they do appear in the shortened succession
found in the xylem of the latter groups, where scalariform and reticulate forms are
usually absent. In Pararaucaria the presence of reticulate tracheids in the succession
is comparable with the similar condition noted by Bailey (1925 : 593) as occurring
sporadically in Araucaria and Agathis, which he has interpreted as intermediate bet-
ween " lower " and " higher " gymnosperms in this respect. Though it is true that
Bailey's observations were made on stem wood and the condition in Pararaucaria is

122 A CONIFEROUS PETRIFIED FOREST IN PATAGONIA

reported from the cone axis only the comparison is worth recording, though fuller
knowledge of developmental sequence in the cone axes of the various groups is
not available for comparison.

The structure of the secondary xylem, in particular the radial pitting of the
tracheid walls and the cross-field pitting of the secondary medullary rays, has often
been used as a comparative basis for assessing relationships in coniferales. The
variability in form and arrangement of tracheid pitting which has been demonstrated
in stem and root wood in individual plants in a variety of families of the Coniferales
is, however, such that the xylem type in the Pararaucaria cones might be included
within the range found, for example, in Pinus, Sequoia, or Araucariaceae (Bailey,
1933 ; Bailey & Faull, 1934 ; Pool, 1929). Cross-field pitting in stem and root
wood is also a variable character (e.g., Bannan, 1944). Cupressoid form of the
pits in the cross-field is found fairly regularly in the Araucariaceae and Cupressaceae,
and it also occurs amongst the Podocarpaceae, Taxodiaceae, Taxaceae, and even
occasionally in the Pinaceae (cf. Phillips, 1941 : 268, 274-277). The condition in
the Araucariaceae and in Taxodium, where the cupressoid cross-field pits may be as
numerous as in Pararaucaria, probably conforms most nearly with that in the fossil
cones. But details of cone-axis wood are available for comparison in very few cases :
in Araucaria the records given by Thomson (1913, pi. 4, fig. 34) are of multiseriate
pitting of the radial tracheid walls of typical araucarian form. This, together
with the very wide pith and narrow ring of xylem (Thomson, 1913 : 4, pi. i, figs.
5, 7) serves readily to distinguish araucarian cone axes from those of Pararaucaria.
No satisfactory direct comparison with cone axis structure of other families of
conifers can be made without further investigation of these. The evidence of the
cross-field pitting taken from stem wood structure would, however, tend to favour
a reference of the fossil cones to relationship with the Taxodiaceae, where too the
same type of tracheidal pitting has been demonstrated in the root of Sequoia semper-
virens (Bailey & Faull, 1934, pi. 103, fig. 29). [See note on p. 138].

The mode of attachment of the vascular supply of the bract and fertile scale
to the axis stele is illustrated in Text-fig. 5, A-D, which is taken from a series of
sections at intervals of approximately 2-5 mm. apart, in the basal half of a cone,
and viewed from the base upwards. It has not been possible to follow out the course
of the bundles from a series through a single leaf-gap : but the vascular supply may
be interpreted from a number of gaps cut at varying levels. The bract supply (b)
comes from the base of the leaf-gap as a single strand, being separate in origin from
the supply of the fertile scale, which is derived as two separate strands (Text-fig. 5A,
fl, fz,) from the sides of the gap above the bract supply ; these two strands subse-
quently fuse to give one large bundle (Text-fig. 5, B-D, /) after each has undergone
torsion through 180°, so that the protoxylem comes to lie on the outer (abaxial)
face of the fusion bundle (/), which is thus inverted alongside the bract supply (b)
in the cortex ; the phloem is not preserved. This is also illustrated in PL 6, figs. 57,
58, (b.s., f.s.). A striking feature, though of subsidiary comparative interest, is
the bulky strand of sclerenchyma which accompanies the inverted vascular supply
of the fertile scale on its adaxial face (PI. 6, figs 57, 58, sc. ; Text-fig. 5, stippling).
This strand, which has its origin in the cortex in the angle above the leaf-gap, forks

A CONIFEROUS PETRIFIED FOREST IN PATAGONIA

123

into two as the fusion bundle passes out of the cortex into the base of the fertile
scale, and the paired strands remain a prominent feature in the fertile scale for about
half its length (Text-fig. 6).

This account of the origin of the bract and fertile scale supplies does not agree
with that of Wieland. He has described, from cut surfaces of cones, the origin of
the bract-fertile scale supply as a " single heavy concentric strand " (1935 : 22, pi. 4,

A

B

rm,

— -b

TEXT-FIG. 5. Pararaucaria patagonica Wieland. Transverse sections of stele of cone
axis from below upwards, to show detachment of bract supply (b) and fertile scale
supply ( /i , /2) , from leaf-gaps. The two traces /i , /2 of the fertile scale supply fuse with
inversion to give a single bundle (/). Xylem black, with protoxylem white; scleren-
chyma of fertile scale in double stippling. V. 30964 c-f. X 6-7.

fig- 3 »' pl. 5> fig 5)- He nevertheless interpreted this condition as being in exact
agreement with that in Pinus, where however, according to the account of Aase
(1915), the bract trace is normally in its origin quite free (except at the cone base)
from the fertile scale supply, as indeed is the case in others of the Pinaceae which
have been examined (e.g., Aase, 1915 ; Radais, 1894). According to the observa-

124 A CONIFEROUS PETRIFIED FOREST IN PATAGONIA

tions on which the present description is based, Paramucaria does resemble the
Pinaceae in this respect, as also the section Bunya of Araucaria, and some of the
Taxodiaceae (Aase, 1915 : 294-7) ; and it has not been possible to confirm Wieland's
observations. In his pi. 4, fig. 3, the supplies to the bract and fertile scale are clearly
separate but in his pi. 5, fig. 5, there is some evidence that a single strand is present
in the cortex outside the leaf-gap : this latter condition, however, is in the cone
base, and such a condition has been noted by Aase (1915 : 282-5, 296) as occurring
in the base of Pinus and Cryptomeria cones, though the normal condition higher up
is to find the bract and fertile scale supplies quite separate in origin.

The external features of the bract and fertile scale at the cone surface have
already been noted : PI. 6, figs. 57-60 and Text-fig. 6 show their structure in series
of tangential longitudinal sections in sequence from inside outwards. The most
conspicuous characters are the thick, persistent and probably woody bract (b),
subtending the thick woody fertile scale (/), which bears on its upper surface a single
large, inverted seed, which is flattened and winged. The bract is free from the
fertile scale for the greater part of its length, and is fused with it only towards the
base (PI. 6, figs. 57-60 and Text-fig. 6). Here the double vascular supply, with the
upper bundle (/.s.) inverted and accompanied by the two strands of sclerenchyma,
is a prominent feature. The lower bundle, which supplies the bract, is single below
but forks about half way up the free part of the bract, and the thick, protruding
tip of the latter is supplied by a number of small vascular strands linked tangentially
by a bulky tissue composed of isodiametric pitted cells, which may be described
as a transfusion tissue (PI. 6, figs. 59, 60, and Text-fig. 6, TT). A curious feature
of the bract as described by Wieland was that it was " pleated " (Wieland, 1935,
pi. 4, figs. 4, 5). His illustrations show this in respect of one flank only of the bracts,
and not in all of them. Closer examination of his figures and of the sections of
the series here described has provided evidence (cf. PL 6, fig. 57) that this apparent
lateral forking or pleating is found only in the bracts cut in inner tangential planes
just outside the region where they are attached to the outer cortex of the axis.
Here they are so close-set that the margins of their bases tend to be laterally con-
fluent where they overlap, so that a kind of reticulum is formed on the surface of
the axis. This is not necessarily a characteristic feature, but one correlated with
the closely imbricate succession of the bracts : it is certainly not one characteristic
of the free distal region of the bract.

The fertile scale (/) is of similar size and thickness to the bract, and is also supplied
in its basal region by a single vascular strand (/.s.) which remains unbranched until
a short distance behind the distal attachment of the seed to the scale. Here it
forks into 3-4 strands, and one of these, a lateral strand, passes up to supply the
seed in the region of the basal foramen (PL 6, fig. 59, and Text-fig. 6F,/.s./.). Above
the insertion of the seed the vascular strands supply the prominent tip of the fertile
scale ; and here too there is a well-marked transfusion tissue as in the bract. The
conspicuous strands of sclerenchyma noted in the base of the fertile scale fade out
about half-way up the scale : their function may have been connected with the
separation of the seed scales at maturity to liberate the seeds. Though the epi-
dermal and hypodermal tissues of the scale in this region are poorly preserved, there

A CONIFEROUS PETRIFIED FOREST IN PATAGONIA

125

is some evidence (PI. 6, fig. 60, and Text-fig. 6n) of a grooved abaxial surface on
the exposed distal face of the scale, possibly a natural feature, and one which has
been noted externally as giving a longitudinally ridged appearance in a few of
the best preserved cones (PI. 6, fig. 63).

H

TEXT-FIG. 6. Pararaucaria patagonica Wieland (cf. PL 6, figs. 57-60). Tangential
longitudinal sections of fertile scales (/) with seed, and bracts (6), taken from base
outwards, (j = transverse section.) Xylem black, sclerenchyma of fertile scale
double stippled, stellate sclerotic wing tissue (w) of testa single stippled, transfusion
tissue (TT) stroked. For. = foramen of seed, fsl — lateral bundle of fertile scale
supplying seed, ts = stony later of testa, Er = radicle of embryo, EC = cotyledons
of embryo, e.s. = ? embryo sac, n — ? nucellus. (A, B, V. 30959 c, e ; C-G, V. 30961 ;
H, i, V. 30960** ; j, V. 309450). x 5.

126 A CONIFEROUS PETRIFIED FOREST IN PATAGONIA

The seeds themselves are large and flattened ; the latter feature appears to be
general and may be a natural character. The usual condition is to find one seed
per scale, inverted and with its basal attachment and vascular supply lateral as
shown in PI. 6, fig. 59 and Text-fig. 6r. Wieland reported one cone only in part
of which he found 2 seeds per scale, but this condition appears to be exceptional
(Wieland, 1935 : 22). All the cones examined appear to be mature ; fully formed
embryos are found preserved in the seeds of some of the cones sectioned, and the
micropyles, directed towards the cone axis, are sealed (PL 7, fig. 73). The seeds
at first sight appear to be embedded in the tissue of the fertile scale on its upper
surface ; but closer examination shows that this is not the case. There is a con-
spicuous " stony " layer in the testa (PL 6, figs. 57-59, Text-fig. 6, t.s.}, consisting
of a deep zone of elongate, thick-walled cells lying with their longer axes at right
angles to the surface. This layer is continuous except for the basal foramen in the
region where the seed is attached. The stony layer is surrounded by an outer
layer, confluent on either side of the seed body with a flattened wing (PL 6, figs.
57-60 and Text-fig. 6, w), which consists of a characteristic lacunar tissue of stellate,
thick-walled cells or sclereids (PL 6, fig. 61), like the " cellules etoilees " of Radais
(1894 : 231) in his description of the wings on the seeds of Abies. This tissue,
resembling an aerenchyma, is markedly different from the tissue of the fertile scale
below, to which it is closely adpressed. It corresponds exactly with the tissue de-
scribed by Radais (1894 : 231, pi. 4, fig. 47) and von Tubeuf (1892 : 205, text-fig.
18) in the wings of the seeds of the Pinaceae, where, in the earlier stages of develop-
ment, the whole of the inner surface of the seed, including the wing, is attached to
the upper surface of the fertile scale, and separation only takes place at a later
stage as the seed matures. The evidence from the specimens of Pammucaria
here examined, which have all been mature, suggests that the seed, including the
wing, is separate from the surface of the scale, except in the region of attachment
at the basal foramen (PL 6, figs. 57-59 and Text-fig. 6). Text-fig. 7 illustrates the
relationship of the seed to the scale in this region : and it can be seen that there is
an " absciss " zone here consisting of an aerenchyma-like tissue continuous into the
foramen, suggesting that at maturity there is rupture of the tissue in this stalk-like
region. This rupture would finally liberate the ripe seed. Without younger stages,
it is hardly possible to decide whether the outermost " wing " of the seed coat is at an
earlier stage attached to the scale, as in the living Pinaceae, or free from it, as in the
living Agathis (where the single seed is median) and some of the Taxodiaceae with
inverted winged seeds (Taiwania, Taxodium}. This point, though a critical one, must
remain undecided. Wieland's specimens were probably also mature and clearly show
the separation of this wing (Wieland, 1935, pi. 4, fig. 5, pi. 5, fig. 2) which he did
not recognize when he interpreted the condition of the seeds as comparable with that
in Araucaria, "inclosed by the drooping curtain-like, expanded, and flattened and
coalesced tip of the seed scale " (p. 22). It was principally on the basis of this feature,
associated with the single seed, that he drew his comparison with the genus Araucaria,
and instituted the name "Pararaucaria" to emphasize the intermediate nature of
the genus; for he interpreted the slender cone axis and narrow pith as pinacean
characters, together with the more or less free bract (p. 23). Florin, however,

A CONIFEROUS PETRIFIED FOREST IN PATAGONIA

127

described Wieland's "curtain-like" structure as the sarcotesta of the seed coat
(1944 : 513), though he did not recognize the seed as winged.

The tissues of the seed inside the stony layer of the testa are poorly preserved in
most specimens : but PI. 6, figs. 57, 58 and Text-fig. 6, c, E, illustrate a cone in

B

TEXT-FIG. 7. Pararaucaria patagonica Wieland. A. Longitudinal section of fertile scale
(/) and bract (b) of cone, cut to one side of median plane to show foramen of seed
(For.) in region of attachment to scale, ts = stony layer of testa, w — wing tissue
of testa : xylem of cone axis and fertile scale shown in black. V. 30966*3. x 6-7.
B. Tissue shown at B in foramen of seed in fig. A. x 240. c. Tissue shown at C in
fig. A. x 240. In B and c, intercellular spaces stippled, thick- walled 1 fibrous cells
in solid black.

which embryos were preserved. The remains of the embryo sac are probably
represented at e.s., and of the nucellus at n. The embryo is seen cut transversely in
the region of the radicle towards the micropylar end of the seed in PL 6, fig. 57. and
Text-fig. 6, c, and in the region of the cotyledons, towards the base of the seed, in
PI. 6, fig. 58, Text-fig. 6, E. There are several (probably eight) cotyledons. PI. 7,
figs. 73, 75 show an embryo in radial longitudinal section in another cone, with the
radicle tip (r) pointing towards the sealed micropyle (m). This is the only specimen
in which it has been possible to demonstrate an embryo in median longitudinal section

128 A CONIFEROUS PETRIFIED FOREST IN PATAGONIA

to confirm the orientation. In Wieland's account (1935 : 22) he refers to a suggestion
of a dicotyledonous embryo, but the preservation of his specimens left this point
inconclusive.

Butts & Buchholz (1940) have shown that in the Pinaceae, polycotyledonary
embryos with cotyledons reaching nine in number are a characteristic feature, while
in the Taxodiaceae, the number varies among the genera from 8 to 2. But in the
remaining families of the conifers embryos with 3 or more cotyledons do occur in
certain genera, though the usual number is 2, so that the character has a limited
diagnostic value.

RELATIONSHIPS

The chief features of Pararaucaria patagonica here described which were not
noted by Wieland (1935) in his account of the species are : the foliation of the
pedicel ; the details of the wood structure ; the detachment of the bract trace
from the cone stele separately from the two traces which form the supply of the
fertile scale ; the winged, detachable nature of the inverted seed and its lateral
position ; and the polycotyledonary embryo.

Wieland's account established the characters of slender cone axis with narrow
pith, and the conspicuous bract scales of the cone bearing in their axils the large
fertile scales each normally with one large inverted seed, which he interpreted as
being enclosed in the scale tissue. On this association of characters he assessed the
systematic position of the genus as intermediate between Pinaceae and Araucaria
(Wieland, 1935 : 23), and accordingly instituted the generic name Pararaucaria.
A re-analysis of the relationships of the genus is attempted in the following table
on the basis of the fuller set of criteria of comparison which can now be used. The
available characters of the petrified cones which might be suitable for giving a
quantitative index of aggregate intergeneric differences are too few in number for
effective use : but the table summarizes the position qualitatively.

Pararaucaria patagonica Wieland, emend.

List of Characters used for Comparison with some other Coniferales,
Numbered for Reference as in Table below.

1. Bract free from fertile scale for greater part of length.

2. Bract large and conspicuous at maturity.

3. Bract trace distinct in origin from traces of fertile scale.

4. Usually one seed per fertile scale.

5. Seeds inverted.

6. Seeds attached laterally.

7. Seeds winged.

8. Seeds free throughout development from fertile scale, except in region of

basal foramen.

9. Several cotyledons (> 4) in embryo of seed.

10. Cone axis slender, with narrow pith in stele.

11. Secondary medullary rays of cone axis wood with pits of cross-field cupressoid

and numerous.

A CONIFEROUS PETRIFIED FOREST IN PATAGONIA 129

TABLE SHOWING POSITIVE CORRELATIONS WITH SOME FAMILIES AND
GENERA OF CONIFERALES.

Characters compared numbered as in list above.
+ = positive correlation in all species described.
© = positive correlation in some species described.
? = doubtful or undescribed character.

Characters of bract, fertile scale and seed. Cone axis

I.

2. 3- 4- 5- 6. 7. i

5. 9. 10.

— •>
n.

+

+ + + + + +

? + • +

+

+ © + + ? ••

+

Pararaucaria

Araucaria

Agathis . - ... + •• + + •• + + 4-

Pinaceae . . .+ •• + •• + + 4- •• +.+

Taxodiaceae . ... + © •• © + © + © . 4- ©

*Romeroites . . ... + ? •• •• ? •• + ?.4- ?

*Cheirolepidaceae .© © ? .. + + •• •• ?.? ?

*Pseudo- Araucaria ... ? ? .. ? + •• •• ?.? ?

* The data for these fossils, which superficially bear some resemblance to Pararaucaria, are taken
from the accounts given by Spegazzini (1924 : Romeroites), Hirmer & H6rhammer (1934 : Cheirolepi-
daceae), Fliche (1895: Pseudo- Araucaria). A revision of the characters in all these types, especially
of the relationship between seed and fertile scale, is necessary before any critical systematic comparison
with Pararaucaria can be made.

It will be seen on summation of positive correlations that the nearest comparison
is with the Taxodiaceae (sensu Pilger, 1926 : 342), which shows the greatest number
of positive correlations. Even allowing for uncertainty on criterion (8) in Para-
aucaria (freedom of seed from scale throughout development), the comparison with
the Taxodiaceae is closer than with the Pinaceae, for on criteria (2) and (n) (large
size of bract ; and cross-field pits numerous, cupressoid) the conditions found in
Pararaucaria and amongst the Taxodiaceae do not characterize the Pinaceae :
and on criteria (i) and (4) (degree of freedom of bract from fertile scale, and number
of seeds per scale), which represent the chief differences between Pararaucaria and
the Taxodiaceae, it may be noted on (i) that in Cryptomeria (and probably also
in the Jurassic genus Elatides) in the Taxodiaceae there is a form intermediate in
respect of degree of fusion of bract and fertile scale, and on (4), that in Taiwania
in the Taxodiaceae a single seed instead of two may sometimes be found, while in
Pararaucaria the two-seeded condition has abnormally been found. In these two
respects Cryptomeria and Taiwania respectively may be regarded as intermediate
between the Pararaucaria condition and a more typical taxodiaceous condition.

Certain features of Pararaucaria have been omitted from the lists above as not
affording sufficiently reliable distinctions : these are the foliation of the pedicel,
the sharp transition from the foliation of the pedicel to the bracts and fertile scales
of the cone itself, the size of the cones and the number of cone-scales per cone, the
radial pitting of the tracheid walls, and the vertical ridging of the fertile scales.
But it may be noted that in each of these features correspondence with characters
occurring amongst the Taxodiaceae may be found. In the case of the radial pitting

GEOL. II, 2. 9

130 A CONIFEROUS PETRIFIED FOREST IN PATAGONIA

of the tracheids, reference may be made to the range of variability in xylem structure
demonstrated in the Redwood, Sequoia sempervirens , by Bailey & Faull (1934,
pi. 103, fig. 29), where the condition in the fossil cones is found in secondary xylem
of root. The vertical ridging of the fertile scale is, so far as I am aware, a feature
characteristic of the Taxodiaceae in the genera Sequoia, Sciadopitys, and Taxodium,
though in the first two of these it is also found to some extent in the bract. Finally,
the general "habit" of the cones is taxodiaceous ; the proportion of bract to fertile
scale, the size of the cones, the vertical ridging of the fertile scale, all combine to
give a habit which on superficial examination one would tend to refer to the
Taxodiaceae.

In the analyses just given, the Taxodiaceae are referred to in the sense of Pilger's
definition (Pilger, 1926 : 347) ; the family in this widely-defined sense shows con-
siderable diversity in certain characters, e.g., the orientation of the seed, and the
number of the cotyledons. The probably heterogeneity of the Taxodiaceae in this
inclusive sense has, however, been generally recognized : Pilger himself separates
the genera into two sub-families and a number of sub-groups, while Hayata (1932)
arranged the genera of Pilger's group in a number of separate families. Florin
has more recently re-examined the relationships of the Taxodiaceae, sensu Pilger,
(Florin, 1931 : 484-491 ; 1940 : 78). He agrees (1931 : 490) with Pilger's classi-
fication, whilst admitting it may be artificial, since it deals with a family inter-
mediate in its general characters between others more sharply defined : but at the
same time he has re-emphasized its diversity and probable heterogeneity. In
particular, he draws attention, amongst other distinctive features, to the character-
istic epidermal characters of the Tasmanian genus, Athrotaxis, which can readily be
used to distinguish it from other genera of Taxodiaceae in the northern hemisphere.

In view of this diversity in the Taxodiaceae, sensu latu, Pararaucaria may mean-
time most conveniently be referred to this family. It most nearly resembles Tai-
-wania in the one-seeded condition, Cryptomeria in the condition of the bract in
relation to fertile scale, and Sequoia and Taxodium in the polycotyledonary embryo,
but resembles no one living genus in its combination of characters. The possibility
remains, however, that the seed and its wing may have been attached, in earlier
stages of development, to the fertile scale as in the Pinaceae : and in this case the
most natural position would be to place Pararaucaria in a separate new family of
Coniferales, intermediate in position between the Taxodiaceae and Pinaceae. In
a recent review by Takhtajan (1953) the phylogeny of the Taxodiaceae has again
been discussed, and he suggests that they probably originated from the earliest and
most primitive Pinaceae, though no fresh evidence is adduced in support of this view.

Wieland (1935 ' 24) referred Pararaucaria to the fossil family Cheirolepidaceae
as defined by Hirmer & Horhammer (1934 : 79) : but this reference was based on
the assumption that the seed was borne in the scale tissue as in the genus Araucaria,
and as has already been demonstrated, this is not the case. Florin (1940 : 36 ;
1944 : 513) suggested that Pararaucaria might be identical with the South American
fossil genus Romeroites, from the province of Neuquen. This genus, as described
by Spegazzini (1924 : 134), is undoubtedly taxodiaceous, but differs from Para-
raucaria chiefly in the thin, many-seeded fertile scales with seeds upright and fleshy,

A CONIFEROUS PETRIFIED FOREST IN PATAGONIA 131

and fusion of bract with fertile scale for the greater part of its length. A more
detailed description of this genus is undoubtedly desirable, but according to Spegaz-
zini's account it does not correspond with Pararaucaria, though the size of the cones
and the general external habit are somewhat similar. Gothan (1950 : 153) in a
very brief note recognized that Pararaucaria had nothing to do with Araucaria,
and accordingly indicated that the genus was misnamed: but he gave no further
description of the cones nor indication of their true relationships, of which, as
has been shown, no more satisfactory definition can be given unless younger stages
in the development of the cones are found.

DISCUSSION

The chief interest of these conifers of the Cerro Cuadrado forest lies in their
relationship with other conifers and their geographical distribution.

Araucaria mirabilis is an extinct species, and appears to have its nearest living
relation in the Queensland " bunya bunya," A. bidwilli, and not in A. araucana or
A. angustifolia, the only living species of Araucaria in South America ; but it might
be considered to represent an ancestral form of these, if the wings of the cone scales
had become reduced in the evolution of the species. This view on the evolution of the
South American living araucarians has already been expressed by Darrow (1936 : 333).
The South American fossil records (cf. Florin's review, 1940 : 33-40) do not include
any which furnish critical evidence of the wingless cone scales of species of the section
Columbea Endlicher, emend. Wilde & Eames : the cone scale from the (?) Upper
Cretaceous of the Cerro Guido in Santa Cruz, which was named Araucarites patagonica
by Kurtz (1902 : 49) and referred to as comparable with the living Araucaria brasilien-
sis, was not figured, and was an incomplete specimen described as representing the
lower part of the scale. Other cone scales in the form of compressions, from Meseta
de Baquero in Santa Cruz, have also been referred to the section Columbea (Feruglio,
1951 : 65). These are more or less incomplete and are not illustrated, but from the
description and from their comparison by Feruglio with Berry's cone scales of Arau-
caria from the nearby Gran Bajo de San Julian (Berry, 1924 : 480, text-figs. 2, 2a)
it appears that they are winged scales. Berry refers to his as " obviously thick and
woody," which suggests that they may have belonged to the section Bunya Wilde &
Eames though they are much smaller than those of the living A. bidwilli. Other
seed scales, which he also compared with Berry's, were described by Feruglio as
Araucaria (Feruglio, 1951 : 39), from the Gran Bajo de San Julian, and they too
might have belonged to the section Bunya. Both Berry's and Feruglio's seed scales
come from localities in the region of the same volcanic complex (the porphyritic
series of the Bahia Laura), which Feruglio has dated as of age mid- Jurassic to
Wealden (1951 : 74), and which also contains the petrified forest around the Cerro
Alto ; but it is unlikely, owing to the incomplete preservation of the compressed
cone scales, that any close comparison can be made between them and the petrified
cone scales of Araucaria mirabilis in the forest, though they are not dissimilar in
size and general form. Feruglio does not believe that the San Julian specimens
are identical with Araucaria mirabilis.

GEOL. II, 2. 9§

132 A CONIFEROUS PETRIFIED FOREST IN PATAGONIA

There is, therefore, as yet no critical record of wingless cone scales in South Ameri-
can fossil floras. The evidence so far provided from sterile shoots with broad flat
leaves might indicate araucarians of the sections Bimya Wilde & Eames, Intermedia
White, or Columbea Endlicher emend. Wilde & Eames ; and petrified woods have
not yet provided reliable evidence for distinction of these sections. The South
American records are, however, relatively scanty, and the absence from them of
critical evidence of fossil species of Columbea, sensu Wilde & Eames, is necessarily
inconclusive for assessment of the age of this section on the South American conti-
nent in comparison with that of species of the section Bunya, of which A . mirabilis
is the only fossil form with critical structural evidence of relationship. Wingless
araucarian cone scales are rare in the fossil records of the Southern Hemisphere as
a whole (Florin, 1940 : 27), but they have been reported from the Jurassic of Australia
and New Zealand (A. grandis Walkom, 1921 : 13, pi, 3, fig. 6 ; Edwards, 1934 : 100),
suggesting that they may have been as ancient as the winged types, which are
abundant in Jurassic rocks, and indicating that they did occur in Mesozoic times
in Australasia, where they are now extinct The evidence, therefore, though scanty,
does not support the attractive hypothesis of the South American evolution of
Columbea ; and accordingly, A . mirabilis probably bears no direct relationship
to the living South American species, but represents an extinct element of an earlier
more widespread araucarian flora, some of the species of which have disappeared.
The presence of such a type as A . mirabilis in South America, however, does provide
another link between the floras of Australasia and South America. A. mirabilis
may well have been a member of the parent stock from which the modern " bunya
bunya " of Queensland sprang, a stock more widely spread than its descendents ; but
in the development of the South American floras it eventually died out, while the
species with wingless cone scales persisted there to the present day. Araucarians
belonging to the Australasian sections Eutacta or Intermedia, with wide, thin, papery
wings on the seed scales, were also represented in earlier South American floras ;
perhaps the best evidence for this is provided by Araucaria pichileufensis , described
by Berry from Rio Pichileufu in Territory of Rio Negro (Berry, 1938 : 59, pi. n,
fig. i). This is a Tertiary species, though its exact age is doubtful (Florin, 1940 :
39), and might suggest that either section Eutacta or Intermedia persisted longer
before extinction on the South American continent than did Bunya, though critical
records are too scanty for such a generalization to be made at this stage. Florin
has renewed the emphasis on the part probably played by Antarctica, whether as
an intercontinental land bridge along which migration could take place, or as the
place of origin of some of the southern genera (Florin, 1940 : 85-6, 92) : and the
distribution of fossil and living species of Araucaria in South America and Austral-
asia supports this hypothesis.

Pararaucaria patagonica represents an extinct genus and presents a different
problem in distribution. The Taxodiaceae, which probably represent the nearest
living relations of this genus, are to-day confined to the northern hemisphere, with
the exception of Athrotaxis, with three species living in Tasmania. The only other
fossil records of this family from the southern hemisphere are three species of Athro-
taxis— A . ungeri (Halle) from the Mesozoic (possibly Lower Cretaceous) of Southern

A CONIFEROUS PETRIFIED FOREST IN PATAGONIA 133

Patagonia, and two other species from the Cretaceous of New Zealand and the Ter-
tiary of Tasmania (Halle, 1913 : 40-44 ; Florin, 1940 : 35, 77) — and Romeroites
argentinensis Speg. (Spegazzini, 1924 : 134-9), from Neuquen, South America
(? Upper Jurassic or Cretaceous). Florin has, however, drawn attention to the
separate and peculiar nature of the genus Athrotaxis, and has suggested that it may
represent a distinct, though probably remotely related, line of descent from the Taxo-
diaceae. Paramucaria adds, then, to the evidence for an earlier more widespread
distribution of taxodiaceous types of plant in the southern hemisphere ; along with
Romeroites, it is now extinct, while Athrotaxis has disappeared from South America.
Pamraucaria is the only genus related to the Taxodiaceae which shows freedom of
bract from fertile scale for the greater part of its length, and so might be regarded
as an earlier member of an evolutionary series of forms leading to some of the modern
Taxodiaceae where fusion is more or less complete. Such an interpretation would
place Pararaucaria in line with that of Florin for the evolution of the coniferalean
ovulate strobilus, where he regards the cohesion of the bract with its axillary seed
scale complex as a condition derivative from more primitive forms where they are
free. However, the Taxodiaceae (sensu Pilger) are an ancient and widespread
family in the northern hemisphere ; among living genera, Sciadopitys is already
known in the Jurassic, and other genera are abundant in younger rocks, especially
in the Tertiary. Hirmer has however (1936 : 65), also included the much older
Upper Permian Pseudovoltzia, the Triassic Voltzia and the Rhaeto-Liassic Sweden-
borgia in this family : in the two former genera, the bract and ovuliferous scale are
more or less free, and in the latter are partially fused. Lack of critical evidence
for the exact geological age of Pararaucaria precludes close phyletic comparison with
other taxodiaceous genera : but as already indicated (p. 102), the most recent
evidence places the age of the petrified forest as at some stage between mid- Jurassic
and Wealden. The characters of Pararaucaria itself certainly support the assignation
of Mesozoic rather than Tertiary age, for in younger floras the majority of the genera
are modern, and Pararaucaria cannot be closely compared with any living genus. A
reference made by Wehrfeld (1935) to wood of palms in the Cerro Cuadrado floras has
never been substantiated ; and the genus Araucaria, as represented by the extinct
A. mirabilis in the same flora, is itself an ancient one, dating back to the Jurassic.
It is probable that Pararaucaria represents an extinct offshoot from some common
earlier stock of the Taxodiaceae, no doubt of northern origin, which underwent
separate evolution south of the equator, as suggested by Florin for the case of Athro-
taxis, rather than a type to be regarded as directly ancestral to any of the living
genera.

The unique one-seeded fertile scale of Pararaucaria is a striking case of homeotic
similarity with the condition found in many other conifers of the southern hemisphere
— Araucaria, Agathis, and the Podocarpaceae, where distribution of the single seed
is associated with the entire woody cone-scale, separate winged seed and entire
cone-scale with fleshy developments respectively. Biologically, the closest com-
parison in this respect with Pararaucaria is with Agathis, where there is also a single
winged seed, but with median insertion on a single cone-scale representing the
completely fused bract and fertile scale. In the living northern Coniferales (ex-

134 A CONIFEROUS PETRIFIED FOREST IN PATAGONIA

eluding Taxales) the cone-scales are normally two to many-seeded, with detachable
seeds, though in the taxodiaceous genus Taiwania, which usually has two winged
seeds per scale, one of these is frequently missing and the condition is similar to
that in Pamraucaria. This modern condition in the northern Coniferales is in
contrast with that amongst the oldest northern conifer floras of the Upper Carboni-
ferous and Permian, where two of the most widespread genera, Lebachia and Ull-
mannia, had one-seeded seed-scale complexes. This condition disappeared relatively
early in the history of northern conifers ; though the Araucariaceae are notable
exceptions during the Mesozoic in northern lands, they too disappeared from northern
floras during early Tertiary times. In any case, this family may have had its
origins in the southern continent and have spread northwards, though final proofs
of this are lacking (Florin, 1940 : 78-82, 90). However, little is known of the early
history of the southern floras ; and in them the one-seeded cone scale may in some
cases represent, as seems probable in Pamraucaria, the result of reduction in number
of seeds on the fertile shoot, rather than the retention of a primitive character.

There is no evidence that the possession of a single seed per seed-scale complex
has conferred any special biological advantage amongst southern conifers, except
in so far as it has made possible, in the genus Araucaria, the distribution of the
single seed still associated with the protective cone-scale tissue in fruit-like fashion,
simulating the samara or nut of angiosperms ; a somewhat similar condition is
found in Podocarpaceae, in association with fleshy developments. In these two
families a condition analogous in some respects with angiospermy has thus been
attained, and the corresponding increase in degree of protection of the seed might
in some measure account for the success of these two families. But in Agathis
and Pararaucaria no such additional protection is associated with the single- seeded
habit, though the larger size of the seeds ensured by the reduction in number may
have had advantages in germination.

CONCLUSIONS

According to this interpretation of the remarkable petrified forest of the Cerro
Cuadrado region, it is unique in a number of respects amongst other petrified forests.
The abundant material of petrified and fertile seed cones of the two dominant types
of tree gives a basis for a fairly satisfactory systematic comparison with other
living and extinct conifers : the structure of the embryos, for example, and of the
vascularization of the cone-scales and ligule in the araucarian cones, are characters
which one can hope to demonstrate only very rarely in fossil conifers, and close
comparison with living species is impracticable without them. Conclusive evidence
of only two species, both based on seed cones, has been found in the forest — Arau-
caria mirabilis ; and Pararaucaria patagonica, whose affinities are probably with
the Taxodiaceae. The detached portions of wood, branches and twigs most probably
belonged to one or other of these species, though critical proof of relationship is
lacking. The presence of the numerous petrified seedlings affords a unique demon-
stration of regeneration, probably of the araucarian species, in the forest. Both
the dominant species are extinct, but one, A. mirabilis, probably disappeared in
the course of evolution of section Bunya of the genus Araucaria, while the other,

A CONIFEROUS PETRIFIED FOREST IN PATAGONIA 135

Paramucaria patagonica, left no descendants, the family of the Taxodiaceae having
disappeared from South America without having played a prominent part in its
floras so far as may be judged from the scanty fossil evidence. The characters of
Pararaucaria in particular suggest an age not younger than Cretaceous for the
forest : for no close taxonomic comparison with any living genus can be made.

My grateful acknowledgments are due to the Dixon Fund of the University of
London for the provision of a grant towards the expenses of section cutting. I
wish also to record my thanks to Mr. W. N. Edwards for his encouragement and
help during the examination of the British Museum collections, and to the Director
of the ficole Superieure de Geologic appliquee in the University of Nancy, to Pro-
fessor R. Florin, Professor T. G. Halle, Professor T. M. Harris and Professor J.
Walton for facilities and help afforded in examining specimens in their care.

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138 A CONIFEROUS PETRIFIED FOREST IN PATAGONIA

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Note for p. 122.

The pith structure in the Pararaucaria cones resembles quite closely that of the
stem in the living Taiwania cryptomerioides. Doyle M. H. & Doyle, J. (1948,
Proc. R. Irish Acad., 52, B : 26, text-fig. 5) have demonstrated that the pith
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genera, though corresponding observations for cone axis anatomy are not available.]

PRESENTED

2 3 NOV 1953

DESCRIPTION OF PLATES
All figures are from untouched photographs.

PI. i, figs, i, 4-6, 8-13 ; PI. 2, figs. 14-20 ; PL 3, figs. 26-37 ; PL 4, figs. 38-42 ; PL 5, fig.
46 ; PL 6, figs. 55, 56, 62-64 '• pl- 7. figs- 74» 75. are taken by reflected light. PL 4, fig. 43
and PL 5, figs. 50-54 are taken from thin sections against a black background.

The photographs in PL i, figs. 3, 7 ; PL 5, figs. 48, 49 ; PL 6, fig. 63, and PL 7, figs. 66-73
were taken by Mr. E. Ashby, Department of Cryptogamic Botany, University of Manchester.

PLATE i

Araucarites sanctaecrucis n. sp. and Coniferous Wood.
FIG. i. Leafy (L) and decorticated (w) branches embedded in matrix. V. 30936. x 0-6.

Coniferous Wood.

FIG. 2. Transverse section of a woody branch, probably 12 years old, showing pith (p],
annual rings in the secondary xylem (x), and remains of cortical tissue (? periderm, c). V.
30937«. x o'8.

FIG. 3. Radial longitudinal section of branch seen in fig. 2, showing pitting on radial walls
of tracheids. V. 309376. x 250.

Araucarites sanctaecrucis n. sp.

FIG. 4. Foliated ? 2-year-old branch, showing lateral branches. V. 30938. x 0-9.

FIG. 5. Detail of abaxial leaf surface from specimen showing grooves and ridges (cf. branch
seen in fig. 4). V. 30939. x 15.

FIG. 6. Partially decorticated ? 3-year-old branch with apparent whorl of ? 2-year-old
lateral branches. V. 30940. x o'8.

Coniferous Wood.
FIG. 7. Detail of pitting shown in fig. 3. V. 309376. x 500.

Araucarites sanctaecrucis n. sp.

FIG. 8. Transversely cut top end of branch shown in figs. 10, n, with (? 4) annual rings
in the secondary xylem (x), and leaf bases (&) and leaf tips (t) transversely cut. V. 30941.

X i.

FIG. 9. Leafy 2-year-old branch with one-year-old branches arranged in two ranks. V.
30942. x 0-9.

FIG. 10. Detail of leaves from the top of the branch shown in figs. 8, n. b — leaf base,
t = leaf tip. V. 30941. X 3-8.

FIG. ii. Leafy branch (? 4-year-old). ¥.30941. X 0-8.

FIG. 12. Defoliated branch (? 5-year-old) showing rhomboidal leaf bases. V. 30943. X 0-9.

FIG. 13. i-year-old leafy branch. V. 30944. x o'8.

Bull. B.M. (N.H.) Geol. 2, 2

PLATE 1

10

13

ARAUCARITES

PLATE 2

Seedlings (cf. Araucaria mirabilis).

FIG. 14. Broken seedling (S), along with separate fragments of branches and twigs,
embedded in matrix. V. 30936. x 0-6.

FIGS. 15-17. Broken seedlings of various shapes (see text). V. 30946—48. x 0-9.

FIG. 18. External view of seedling showing periderm (pd) and woody core (x). V. 30949.
X 0-9.

FIG. 19. Median longitudinally-cut surface of half seedling shown in fig. 18. x 0-9.

FIG. 20. Transversely-cut surface of swollen region of seedling similar to those shown in
figs. 17, 18. V. 30950. x 0-9.

FIG. 21. Radial longitudinal section of top end of seedling similar to that shown in fig. 15.
p = pith. V. 30951^. x o'8.

FIG. 22. Transverse section of central swollen region of seedling similar to those shown in
figs. 17-20. pd = periderm, x = secondary xylem. V. 3095205. x o'8.

FIGS. 23-25. Transverse sections through top, middle and basal end respectively of seedling
similar to that shown in fig. 15. p = pith, pr — primary xylem, x = secondary xylem (cf.
Text-fig, i). V. 3og$ib-d. x 25.

Bull. B.M. (N.H.) Geol. 2, 2

PLATE 2

14

20

Araucarian Seedlings

PLATE 3

Araucaria mirabilis (Spegazzini).

FIG. 26. Petrified cones embedded in matrix. V. 30953. x 0-4

FIG. 27. Cone broken transversely half way up axis, showing pith (p), vascular bundles (v),
and cone-scales with seeds (s). V. 39054. x 0-9.

FIG. 28. External features of cone shown in fig. 27. / = ligule, a = apophysis of bract.
X 0-9.

FIG. 29. Detail of external features of cone-scales shown in fig. 28. / = ligule, a = apo-
physis of bract, w — wing. x 4-1.

FIG. 30. Base of cone shown in fig. 35 ; showing transversely cut pedicel (pi), and gradual
transition from sterile to fertile scales at base of axis. V. 30955. x 0-8.

FIG. 31. Small cone, showing overlapping laminar tips of bracts. V. 30956. x 0-8.

FIG. 32. Cone shown in fig. 31, median longitudinal surface, showing wide pith (p) and
immature or sterile cone-scales (c). x 0*8.

FIG. 33. Detail of cone scales in fig. 32, showing ligule (/) and bract (b) ; no ovules have
been formed. x 4*1.

FIG. 34. Detail of fig. 31, showing laminar tips of bracts. x 4-1.

FIG. 35. Polished longitudinally cut surface of half cone shown in fig. 30. A = cone-scale
shown in detail in figs. 36-37 ; B = cone-scale cut in approximately median longitudinal
section ; E = embryo in seed, x 0-9.

FIG. 36. Detail of fig. 35 at A. / = ligule, a = apophysis of bract (artificially split),
S = seed (cut to one side of median plane), l.s. — ligular sulcus. x 4-1.

FIG. 37. Surface view of half cone-scale shown in section in fig. 36. / — ligule, a = apo-
physis of bract, x 4-1.

Bull. B.M. (N.H.) Geol 2,

PLATE 3

•s*4>p

~^:-^*

ARAUCARIA MJRABILIS

PLATE 4
Araucaria mirabilis (Spegazzini).

FIG. 38. Half cone in surface view, showing persistent laminar tips of bracts of cone-scales.
Structure illustrated in figs. 39-45, and PI. 5, fig. 54. V. 30957. x 0-8.

FIG. 39. Detail of fig. 38, showing persistent laminar tips of bracts (t), with longitudinal
striation. x 4-1.

FIG. 40. Detail of median longitudinally cut face at mid left of half cone in fig. 42, with
apophysis of bract (a), persistent laminar tip of bract (t), ligule (/), embryo (e), (A = ? absciss
zone.) x 4*1.

FIG. 41. Detail of embryo from fig. 42, showing two cotyledons, x 4'i.

FIG. 42. Polished median face of half cone shown in fig. 38 ; p = pith, v.b. = vascular
bundles of axis ; details of cone scales in figs. 40, 41. x 0-8.

FIG. 43. Tangential longitudinal section of cone (cf. figs. 38-42), taken at level of per-
sistent laminar tip of bract, a = apophysis of bract, t = laminar tip, v.b.b. = vascular bundles
of bract. V. 30957. x 4-1.

FIG. 44. Detail of vascular strand in ligule. tr = spiral tracheids. V. 30957. x 208.

FIG. 45. Detail of vascular bundle from persistent laminar tip of bract shown in fig. 43.
d = ? resin duct, v.b. = vascular bundle, x 55.

Bull. B.M. (N.H.) Geol. 2, 2

PLATE 4

42

45

AKAUCARIA MIRABILIS

PLATE 5

Araucaria mirabilis (Spegazzini).

FIG. 46. Surface view of mature seed cone whose structure is illustrated in figs. 47-53.
/ = ligule, a — apophysis of bract. V. 30958. x 4-1.

FIG. 47. Detail of tangential longitudinal section of cone-scale from fig. 50 ; T.S. = testa
of seed (stony layer), F = surface tissue of fertile scale, en = endosperm, E = embryo. V.
309580. x 13.

FIG. 48. Detail of embryo seen obliquely cut in transverse section of cone in fig. 51 (E) ;
co = cotyledon, en — endosperm. V. 309586. x 25.

FIG. 49. Transverse section of embryo from tangential longitudinal section of cone (cf.
fig. 47), showing two cotyledons (co). V. 30958$. x 25.

FIG. 50. Tangential longitudinal section of cone, showing cone-scales with seeds, c = cone-
scale with wing, S = seed with endosperm and embryo, F = surface tissue of fertile scale. V.
309580. x 4-1.

FIG. 51. Transverse section of cone ; T.S. = testa of seed (stony layer), en = endosperm,
E = embryo, ss = suspensors. V. 30986. x 4*1.

FIG. 52. Radial longitudinal section of cone ; lettering as in fig. 51. V. 3O958c. x 4*1.

FIG. 53. Tangential longitudinal section of cone, taken at i mm. from outer surface ;
/ = ligule with vascular bundles, b = bract with vascular bundles. V. 30958^. x 8. ,

FIG. 54. Tangential longitudinal section of cone illustrated in PI. 4, taken at level of ligule
(/) and showing vascular bundles of ligule (v.b.l.), and vascular bundles of bract (v.b.b.) with
? resiri ducts (d). V. 30957$. x 8.

Bull. B.M. (N.H.) Qeol. 2, 2

PLATE 5

54

ARAUCARIA MIRABILIS

PLATE 6

Pavaraucaria patagonica Wieland.

FIG. 55. Cone showing fertile scales (/) and bracts (6), the surface features weathered.
V. 30959. x i.

FIG. 56. Cone showing less marked degree of weathering. V. 30960. x i.

FIGS. 57-60. Tangential longitudinal sections (cf. also Text-fig. 6) through fertile scales
and bracts, taken from base outwards. / = fertile scale, f.s. — vascular supply of fertile
scale, f.s. I. — lateral bundle supplying seed in region of basal foramen, sc = sclerenchyma of
fertile scale, b = bract, b.s. — vascular supply of bract, T.T. = transfusion tissue, w = wing
tissue of testa of seed (cf. fig. 61), t.s. = stony layer of testa of seed, F = basal foramen of seed,
E.r. = radicle of embryo, E.G. = cotyledons of embryo, n — nucellus, e.s. = embryo sac.
Figs. 57-59, V. 3096ia-c. Fig. 60, V. 30960^ x 8.

FIG. 61. Stellate sclerotic tissue of wing of seed (cf. fig. 59, w). t.s. = stony layer of testa
of seed. V. 30961^. x 50.

FIG. 62. Cone with surface features much weathered, showing foliated pedicel (pi.).
V. 30962. x i.

FIG. 63. Cone showing surface features of fertile scales and bracts, the fertile scales with
vertical grooving of protruding abaxial surface. V. 30963. x 0-8.

FIG. 64. Detail of leaves on pedicel from cone in fig. 62, showing longitudinally striated
abaxial surface. V. 30962. x 3-3.

Bull. B.M. (N.H.) Geol. 2, 2

PLATE 6

K v- j^»f^ '«V;V

60

PARARAUCARIA PAT AGON 1C A

PLATE 7
Pararaucaria patagonica Wieland.

FIG. 65. Transverse section of stele of cone axis in lower half of cone, showing narrow
pith (p) and thick cylinder of xylem (x), with bract trace at b. V. 30964^. x 11-7.

FIGS. 66-72 illustrate structure of xylem of cone axis. V. 30959^.

FIG. 66. Radial longitudinal section of transitional region of xylem, showing from right to
left the sequence from protoxylem (px) through metaxylem (mx) to secondary xylem. x 166.

FIG. 67. Detail of pitting in primary xylem from fig. 66, showing spiral pitted tracheids
(s.p.) and reticulate tracheids (ret.). x 417.

FIG. 68. Radial longitudinal section of xylem showing, on the right, reticulate tracheids
of primary xylem, and on the left, the radial walls of secondary tracheids (sec.) with uniseriate
bordered pits. m.r. = medullary ray. x 166.

FIG. 69. Detail from fig. 68, showing pits on radial walls of secondary xylem tracheids, and
part of medullary ray (m.r.) with cross-field pitting, x 333.

FIG. 70. Radial longitudinal section of secondary xylem at base of cone axis, showing
uniseriate and biseriate (bi) radial pitting of tracheids, and medullary ray with cross-field pitting
(m.r.) x 166.

FIG. 71. Detail of biseriate pitting on radial walls of secondary tracheids, from fig. 70.

x 333-

FIG. 72. Radial longitudinal section of secondary xylem in region of medullary ray, showing
cupressoid cross-field pitting (c). (Longitudinal axis of cone runs across the photo.) x 417.

FIG. 73. Radial longitudinal section of fertile scale with seed and embryo, seen (reversed)
at E in fig. 75, showing orientation of embryo, c = cotyledons, r = tip of radicle, m = closed
micropyle. V. 309650 x 8.

FIG. 74. External features of long cone, showing weathered surfaces of fertile scales (/) and
bracts (b). V. 30965. x 1-5.

FIG. 75. Median longitudinally cut surface of cone in fig. 74, showing fertile scales bearing
seeds with embryos (E). 5 = axis stele. x 1-5.

Bull. B.M. (N.H.) Geol. 2, 2

PLATE 7

PA RA RA UCA RJA PA TA GONICA

PRESENTED

2 6 NOV 1953

PRINTED IN GREAT BRITAIN BY
A D L A R D AND SON, LIMITED
BARTHOLOMEW PRESS, DORKING

. _____ . Nov 1g53

'HE SOLUTION OF
THE PILTDOWN PROBLEM

J. S. WEINER, K. P. OAKLEY

AND

W. E. LE GROS CLARK

BULLETIN OF
THE BRITISH MUSEUM (NATURAL HISTORY)

GEOLOGY Vol. 2 No. 3

LONDON: 1953

BULLETIN OF
THE BRITISH MUSEUM (NATURAL HISTORY)

GEOLOGY

The following papers appeared in Volume I (1949-52) :

Price
No. I (1949). The Pterobranch Rhabdopleura in the English Eocene.

H. D. Thomas & A. G. Davis js. 6d.

No. 2 (1949). A Reconsideration of the Galley Hill Skeleton. K. P.

Oakley & M. F. Ashley Montagu ...... ^s.

No. 3 (1950). The Vertebrate Faunas of the Lower Old Red Sandstone

of the Welsh Borders. E. I. White.

Pteraspis leathensis White a Dittonian Zone-Fossil. E. I.

White j$. 6d.

No. 4 (1950). A New Tithonian Ammonoid Fauna from Kurdistan,

Northern Iraq. L. F. Spath ....... IDS.

No. 5 (1951). Cretaceous and Eocene Peduncles of the Cirripede Euscal-

pellum. T. H. Withers 5$.

No. 6 (1951). Some Jurassic and Cretaceous Crabs (Prosoponidae).

T. H. Withers 55.

No. 7 (1952). A New Trochiliscus (Charophyta) from the Downtonian

of Podolia. W. N. Croft IDS.

No. 8 (1952). Cretaceous and Tertiary Foraminifera from the Middle

East. T. F. Grimsdale ........ IDS.

No. 9 (1952). Australian Arthrodires. E. I. White .... 155.

No. 10 (1952). Cyclopygid Trilobites from Girvan. W. F. Whittard . 65.

THE SOLUTION OF
THE PILTDOWN PROBLEM

BY

J. S. WEINER

Department of Anatomy, University of Oxford

K. P. OAKLEY

Department of Geology, British Museum (Natural History)

W. E. LE GROS CLARK, F.R.S.

Department of Anatomy, University of Oxford

Pp. 139-146 ; Pis. 8, 9

BULLETIN OF
THE BRITISH MUSEUM (NATURAL HISTORY)

GEOLOGY Vol. 2 No. 3

LONDON: 1953

THE BULLETIN OF THE BRITISH MUSEUM
(NATURAL HISTORY), instituted in 1949, is
issued in five series corresponding to the Departments
of the Museum, and, an Historical Series.

Parts will appear at irregular intervals as they
become ready. Volumes will contain about three or
four hundred pages, and will not necessarily be
completed within one calendar year.

This paper is Vol. 2, No. 3 of the Geological series.

PRINTED BY ORDER OF THE TRUSTEES OF
THE BRITISH MUSEUM

Issued November zist, 1953 Price Three Shillings and Sixpence

THE SOLUTION OF
THE PILTDOWN PROBLEM

By J. S. WEINER, K. P. OAKLEY & W. E. LE GROS CLARK

NOTE. — The curator of a palaeontological collection, which may contain rare specimens of
great scientific importance, is frequently faced with the problem of whether to allow such
specimens to be reinvestigated by treatment with acids, sectioning, removal of fragments for
chemical analysis, or other methods which might seem to involve damage to a unique object.
The cautious attitude of a previous generation has undoubtedly preserved for their successors
many fossils which, for example, might have been damaged by mechanical treatment in the
past, but can now be developed in perfection by more recently devised chemical methods.
In the case of the Piltdown relics, one can be certain that after they came into the late Sir
Arthur Smith Woodward's possession they would not have been treated or tampered with
either chemically or physically. The decision to submit these specimens to a rigorous re-
examination was made some years ago ; the final result, unexpected at the time, will be found
in the following pages.

W. N. EDWARDS, Keeper of Geology.

SINCE the report, some forty years ago,1 of the discovery of several cranial fragments,
a portion of a mandible and a canine tooth at Piltdown in Sussex, the problem of
the " Piltdown skull " has been the subject of continuous controversy. Some
authorities have accepted all the remains as those of an extinct type of hominid.
But it is probably true to say that most anthropologists have remained sceptical
or frankly puzzled by the contradictions which they present, for (apart altogether
from other details) the combination of a cranium closely similar to that of Homo
sapiens with a mandible and canine tooth of simian form seemed too incongruous.
It has been suggested, indeed, that they really represent the fortuitous association
of a Pleistocene human cranium with the remains of a fossil ape which had perhaps
been secondarily derived from an earlier geological deposit. The application of
the fluorine test (Oakley & Hoskins, 1950), however, made it quite clear that the
mandible and canine were certainly not older than the cranium and, on the assumption
that they were all genuine fossils, it naturally appeared to lend some support to those
who held them to be contemporaneous and to belong to the same individual. It
is also a fact that the remarkably flat wear of the molar teeth in the mandibular
fragment is quite unlike that normally found in apes at a corresponding stage of
dental attrition (though similar to the type of wear characteristic of the hominid
dentition), while the mode of wear of the large canine tooth is also different from
that which occurs in apes. But there is another possible explanation of the apparent
contradictions shown by the Piltdown remains : that the mandible and canine
tooth are actually those of a modern ape (chimpanzee or orang) which have been
deliberately faked to simulate fossil specimens. It was not till one of us (J. S. W.)
in the course of personal discussions put forward this proposition fairly and squarely
as the only possible solution of the Piltdown puzzle, pointed out that the organic
content of the mandible had never been examined, and moreover demonstrated
experimentally that artificial abrasion of the teeth of a chimpanzee combined with
appropriate staining produced an appearance astonishingly similar to the Piltdown

1 Dawson & Woodward, 1913.
GEOL. ii, 3. 10

142 THE SOLUTION OF THE PILTDOWN PROBLEM

molars and canine, that we decided on a critical re-study of all the Piltdown
material with this specific possibility directly in view. The results of our in-
vestigations have now demonstrated quite clearly that the mandible and canine
are indeed deliberate fakes. The evidence1 for this conclusion is briefly as follows :

EVIDENCE OF THE ARTIFICIAL ABRASION OF THE
PILTDOWN TEETH

Molar Teeth of the Mandible

(1) The occlusal surfaces (particularly of M2) are planed down over almost their
whole extent to a flatness which is much more even than that normally produced
by natural wear (PI. 9, fig. 2).

(2) The borders of the flat occlusal surfaces — particularly the lateral borders — are
sharp-cut and show no evidence of the bevelling which is usually produced by
natural wear (PI. 9, fig. i).

(3) The centre of the talonid basin in M2 is unworn, and is bounded by a sharp-cut
and unbe veiled border of the planed surface of the crown. This appearance would
be produced by artificial abrasion but would not be expected in natural wear (PL 9,
fig. i).

(4) The surface of the areas of dentine exposed on the antero-medial cusps of the
two molars is quite flat and flush with the surrounding enamel, instead of forming
a depression as would be expected in natural wear.

(5) In both molars much more dentine has been exposed on the antero-internal
than the antero-external cusps. But in the course of natural attrition the lateral
cusps of lower molar teeth are normally worn down more rapidly (and thus usually
show a greater exposure of dentine) than the medial cusps (PI. 9, fig. i).

(6) The degree of wear in the two molars, Mx and M2, is almost identical. But in
early stages of natural attrition Mt is commonly (though not always) more severely
worn than M2 (PI. 9, fig. 3).

(7) The planes of the flat occlusal surfaces of the two molars are not congruous,
i.e., they do not fit together to form a uniform contour. Unless the teeth have been
displaced from their natural position after death (for which there is no evidence),
this incongruity is difficult to explain by natural wear (PI. 8; 9, fig. 3).

(8) Inspection of the isolated molar tooth (referred to the specimen called Pilt-
down II) with a binocular microscope shows that the occlusal surface of the enamel
has been finely scratched, as though by an abrasive.

Canine Tooth

(1) The mode of wear of this tooth is unlike that found normally either in ape
or human canines, for the abraded surface has exposed the dentine over the entire
lingual surface from medial to distal border and at one point actually reaches the
apex of the pulp cavity (Pi. 9, fig. 5).

(2) The condition of the apex of the root, and the wide and open pulp cavity seen
in an X-ray photograph, indicate fairly certainly that the canine was still incom-
pletely erupted or had only just recently completed its eruption. But this would
be incompatible with the severe attrition of the crown if the latter were naturally
produced (PI. 9, fig. 4).

1 The full evidence will be discussed in detail in a later number of this Bulletin.

THE SOLUTION OF THE PILTDOWN PROBLEM 143

(3) X-ray examination shows no evidence of the deposition of secondary dentine
(with a constriction of the pulp cavity) which might be expected if the severe abrasion
of the lingual surface of the crown were the result of natural attrition (PL 9, fig. 4).

(4) The abraded surface of the crown shows fine vertically disposed scratches
(as seen under a binocular microscope) which suggest the application of an abrasive
(PI. 9> %. 5).

EVIDENCE OF THE FLUORINE CONTENT

The fluorine method as applied in 1949 (and reported in full in 1950) served well
enough to establish that neither the Piltdown cranium nor the mandible was Lower
Pleistocene. It did not distinguish (nor at that time was it intended to distinguish)
Upper Pleistocene from later material. The rate of fluorination at this site has prob-
ably not been high enough to give a clear separation between Upper Pleistocene and,
say, Early Post-glacial bones. Moreover the method of analysis used in 1949 was
accurate only within rather wide limits when applied to samples weighing less than
10 milligrams, with the consequence that even the difference between the fluorine
contents of fossil and modern specimens was obscured where the samples were of
that order of magnitude. Improvements in technique have since led to greater
accuracy in estimating small amounts of fluorine, and it therefore seemed worth
while submitting further samples of the critical Piltdown specimens for analysis
in the Government Laboratory. The new estimations, based mainly on larger
samples, were made by Mr. C. F. M. Fryd. The following summary of the results
leaves no doubt that, whereas the Piltdown cranium may well be Upper Pleistocene
as claimed in 1950, the mandible, canine tooth and isolated molar are quite modern.

%F

o/ p ~V~P~Q :

/o^ /Of2u5

Minimum F-content of local U. Pleistocene bones . 0*1 0-4

Ditto. Upper Pleistocene teeth 1 . . . . o • i 0-4

Piltdown cranium I. . . . . . o • i 0-8

Piltdowncranium.il: frontal . . . . o-i 0-8

Piltdown cranium II : occipital . . . . 0-03 0*2

Piltdown mandible (bone) . . . i . <o-o3 <o-2

Molar of Piltdown mandible . . . . <o-o4 <o-2

Piltdown canine ...... <o-o3 <o-2

Isolated molar (Piltdown II) .... <o-oi <o-i

Molar of Recent chimpanzee . . . . <o-o6 <o*3

1 All the tooth samples were dentine.

EVIDENCE OF THE ORGANIC CONTENT

To regard the organic content of bones and teeth as a measure of their antiquity
has long been regarded as fallacious, and for that reason no serious attempt has
ever been made to test the relative ages of the various Piltdown specimens by that
means. However, extensive chemical studies of bones from early occupation sites in
North America by Cook & Heizer (1947) have shown that in bones preserved under
broadly the same conditions the nitrogen of their protein (ossein) is lost at a relatively
slow, and on an average almost uniformly declining, rate. Thus, N-analysis, used
with discretion, can be an important supplement to F-analysis, and also for the
relative dating of specimens too recent to be within the range of the fluorine method.

I44 THE SOLUTION OF THE PILTDOWN PROBLEM

Dr. J. D. H. Wiseman and Mrs. A. Foster in the Department of Minerals of the British
Museum have devised a method of estimating very small quantities of nitrogen, and
Mrs. Foster, using this new method, determined the nitrogen content of a series of
samples of the Piltdown material and of selected controls. The following is a
summary of the results of this work which agree with all the other evidence
indicating that the Piltdown mandible, canine and isolated molar (II) are modern.
(The possibility that the Piltdown specimens were steeped in a gelatinous preserv-
ative has been borne in mind ; if this had been the explanation of their
nitrogen-content, the cranial bones which are porous would have shown more
nitrogen than the highly compact dentine of the teeth ; whereas the reverse is true.)

Nitrogen-content of Bone Samples

%N
Fresh bone . . . . . . .4-1

Piltdown mandible . . . . . 3 • 9

Neolithic bone (Kent) . . . . i • 9

Piltdown I cranial bones (average) . . .1-4
Piltdown II frontal . . . . . i • i

Piltdown II occipital . . . . .0-6

U. Pleistocene bone (London) . . .0-7

Nitrogen-content of Dentine Samples

%N
Chimpanzee molar . . . . .3-2

Piltdown canine . . . . . .5-1

Piltdown I molar . . . . . . 4 • 3

Piltdown II molar . . . . .4-2

U. Pleistocene equine molar (Piltdown) . .1-2
U. Pleistocene human molar (Surrey) . .0-3

EVIDENCE OF THE COLOURING OF THE PILTDOWN SPECIMENS

A black coating — ferruginous according to Dawson & Woodward (1914: 87)—
covers most of the surface of the Piltdown canine. When this tooth and the molars
were sampled in 1948, one of us (K. P. O.) noted that " below an extremely thin
ferruginous surface stain the dentine was pure white, apparently no more altered
than the dentine of Recent teeth from the soil." Examination by Dr. G. F. Claring-
bull in the Department of Minerals has now shown that the coating on the canine
is in fact non-metallic, it is a tough, flexible paint-like substance, insoluble in the
common organic solvents, and with only a small ash-content. The extreme white-
ness of the dentine and the nature of the black skin are thus both consistent with
the evidence presented above for the essential modernity of the canine.

The mandible is of a reddish-brown colour which, though rather patchy, matches
closely enough that of the cranial fragments to raise no suspicion that all the remains
(from the original Piltdown site) might not belong to one skull. The frontal frag-
ment stated to have been found at a second site (Piltdown II) is also of a similar
brown colour but differs noticeably from the darker greyish-brown occipital frag-
ment from the same site. That the colour of all these fragments is due to iron
oxides has been confirmed by direct analysis in the Government Laboratory.

THE SOLUTION OF THE PILTDOWN PROBLEM 145

But whereas the cranial fragments are all deeply stained (up to 8% of iron) through-
out their thickness, the iron staining of the mandible is quite superficial. A small
surface sample analysed in 1949 contained 7% iron, but, when in the course of our
re-examination this bone was drilled more deeply, the sample obtained was lighter
in colour and contained only 2-3% of iron. The difference in iron staining is thus
also in keeping with the other evidence that the jaw and the cranium are not
naturally associated.

Smith Woodward recorded (1948 : 59 ; see also 1935 : 134) that " the colour of
the pieces which were first discovered was altered a little by Mr. Dawson when he
dipped them in a solution of bichromate of potash in the mistaken idea that this
would harden them." Direct chemical analysis carried out by Drs. M. H. Hey and
A. A. Moss in the Department of Minerals at the British Museum (Natural History),
as well as the X-ray spectrographic method of Mr. E. T. Hall in the Clarendon
Laboratory, Oxford University, confirmed that air the cranial fragments seen by
Smith Woodward in the spring of 1912 (before he commenced systematic excavations)
do contain chromate ; on the other hand, there is no chromate in the cranial frag-
ments subsequently collected that summer — either in the right parietal, or in
the small occipital fragment found in situ by Smith Woodward himself. This being
so, it is not to be expected that the mandible (which was excavated later and in
the presence of Smith Woodward, 1948 : u) would be chromate stained. In fact, as
shown by direct chemical analysis carried out in the Department of Minerals of
the British Museum, the jaw does contain chromate. It is clear from Smith
Woodward's statement about the staining of the cranial fragments of Piltdown I
(which we have verified), that a chromate staining of the jaw could hardly have
been carried out without his knowledge after excavation. The iron and chromate
staining of the Piltdown jaw seems to us to be explicable only as a necessary part
of the deliberate matching of the jaw of a modern ape with the mineralized cranial
fragments.

This grave interpretation, which we have found difficult to avoid, receives
support from the finding that the frontal and occipital fragments labelled Piltdown
II (and found three years later) contain small amounts of chromate. The piece of
frontal bone, anatomically, could form part of the cranium of Piltdown I, and in
colour and in its content of nitrogen and flourine it resembles the first occipital of
Piltdown I rather than that of Piltdown II. Just as the isolated molar almost
certainly comes from the Piltdown mandible, it seems only too likely that this frontal
fragment originally belonged to the cranium of Piltdown I.

From the evidence which we have obtained, it is now clear that the distinguished
palaeontologists and archaeologists who took part in the excavations at Piltdown
were the victims of a most elaborate and carefully prepared hoax. Let it be said,
however, in exoneration of those who have assumed the Piltdown fragments to
belong to a single individual, or who, having examined the original specimens, either
regarded the mandible and canine as those of a fossil ape or else assumed (tacitly
or explicitly) that the problem was not capable of solution on the available evidence,
that the faking of the mandible and canine is so extraordinarily skilful, and the
perpetration of the hoax appears to have been so entirely unscrupulous and inex-
plicable, as to find no parallel in the history of palaeontological discovery.

Lastly, it may be pointed out that the elimination of the Piltdown jaw and teeth
from any further consideration clarifies very considerably the problem of human

146 THE SOLUTION OF THE PILTDOWN PROBLEM

evolution. For it has to be realized that " Piltdown Man " (Eoanthropus) was
actually a most awkward and perplexing element in the fossil record of the Homi-
nidae, being entirely out of conformity both in its strange mixture of morphological
characters and its time sequence with all the palaeontological evidence of human
evolution available from other parts of the world.

REFERENCES

COOK, S. F., & HEIZER, R. F. 1947. The quantitative investigation of aboriginal sites :

Analyses of human bone. Amer. J. Phys. Anthrop., Washington (n.s.) 5: 201-220.
DAWSON, C., & WOODWARD, A. S. 1913. On the Discovery of a Palaeolithic Human Skull

and Mandible in a Flint-bearing Gravel overlying the Wealden (Hastings Beds) at Piltdown,

Fletching (Sussex). Quart. J. Geol. Soc. Lond., 69: 117-144.
1914. Supplementary note on the Discovery of a Palaeolithic Skull and Mandible

at Piltdown (Sussex). Quart. J. Geol. Soc. Lond., 70 : 82-93.
OAKLEY, K. P., & HOSKINS, C. R. 1950. New Evidence on the Antiquity of Piltdown Man.

Nature, Lond., 165 : 379-382.
WOODWARD, A. S. 1935. Recent Progress in the study of Early Man. Rep. Brit. Ass.,

London, 105 : 129-142.
1948. The Earliest Englishman. 118 pp., 3 pis. London.

EXPLANATION OF PLATES

PLATE 8

FIG. i. The Piltdown mandible. Natural size, (i) Outer view ; (2) inner view.

[Photographs by C. Norton.]

PLATE 9

FIG. i. The second molar tooth of the Piltdown mandible viewed from its occlusal aspect.
X 4 diam. approx. Note the relatively large area of dentine exposed on the antero-internal
cusp, and the sharp margin separating the central depression (talonid basin) of the tooth from
the completely flattened cusps. Note also the sharp external margin of the " occlusal " surface,
with no bevelling.

FIG. 2. The second molar tooth of the Piltdown mandible, viewed from the medial aspect.
X 4 diam. approx. Note the extreme evenness of the flat " occlusal " surface.

FIG. 3. The first and second molar teeth of the Piltdown mandible viewed from behind.
X 4 diam. approx. Note that the " occlusal " planes of the two teeth are set at an angle to
each other.

FIG. 4. Radiograph of the Piltdown canine taken in an approximately antero-posterior
plane. X 2 diam. approx. Note the thinness of the dentine on the lingual surface towards
the apex of the tooth. At one point here the pulp cavity appears to have been exposed and
the opening plugged with some material containing radio-opaque particles. The pulp cavity
has been packed from the other end with mineral grains.

FIG. 5. The Piltdown canine tooth viewed from the lingual aspect, x 4 diam. approx.
Note the fine scratches disposed mainly in a vertical direction. A little above the middle of
the surface, towards the apex of the tooth, is seen a small oval area of lighter shade, marking
the point where the pulp cavity has been opened by abrasion.

FIG. 6a-c. Samples of bone, drilled by L. E. Parsons using a parallel burr size 6, from :
(a) Piltdown mandible ; (b) mandible of Recent chimpanzee ; (c) Piltdown skull (right parietal).
X 15. Note that a and b consist of minute shavings, whereas c is a powder.

[Photographs : C. H or ton ; X-ray of Fig. 4 by P. E. Purves.}

D

Bull. B.M. (N.H.) Geol. 2, 3

PLATE

THE PILTDOWN MANDIBLE.

Bull. B.M. (N.H.) Geol. 2,

PLATE g

6a

6b

6c

PILTDOWN TEETH AND BONE.

PRESENTED

2 1 NOV 1953

PRINTED IN GREAT BRITAIN BY
ADLARD AND SON, LIMITED
BARTHOLOMEW PRESS, DORKING

1 6

SOME UPPER CRETACEOUS

AND EOCENE FRUI ^S

FROM EGYPT

M. E. J. CHANDLER

BULLETIN OF
THE BRITISH MUSEUM (NATURAL HISTORY)

GEOLOGY Vol. 2 No. 4

LONDON : 1954

BULLETIN OF
THE BRITISH MUSEUM (NATURAL HISTORY)

GEOLOGY

The following papers appeared in Volume I (1949-52) :

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No. I (1949). The Pterobranch Rhabdopleura in the English Eocene.

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No. 2 (1949). A Reconsideration of the Galley Hill Skeleton. K. P.

Oakley & M. F. Ashley Montagu ...... 55.

No. 3 (1950). The Vertebrate Faunas of the Lower Old Red Sandstone

of the Welsh Borders. E. I. White.

Pteraspis leathensis White a Dittonian Zone-Fossil. E. I.

White js. (>d.

No. 4 (1950). A New Tithonian Ammonoid Fauna from Kurdistan,

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pellum. T. H. Withers 55.

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No. 10 (1952). Cyclopygid Trilobites from Girvan. W. F. Whittard . 6s.

SOME UPPER CRETACEOUS AND EOCENE
FRUITS FROM EGYPT

BY

MARJORIE E. J. CHANDLER

(With Appendices by M. Y. HASSAN and M. I. YOUSSEF)

Pp. 147-187 ; Pis. 10-16 ; i Text-figure

BULLETIN OF
THE BRITISH MUSEUM (NATURAL HISTORY)

GEOLOGY Vol. 2 No. 4

LONDON: 1954

THE BULLETIN OF THE BRITISH MUSEUM
(NATURAL HISTORY), instituted in 1949, is
issued in five series, corresponding to the Departments
of the Museum, and an Historical series.

Parts will appear at irregular intervals as they
become ready. Volumes will contain about three or
four hundred pages, and will not necessarily be
completed within one calendar year.

This paper is Vol. 2, No. 4 of the Geological series.

PRINTED BY ORDER OF THE TRUSTEES OF
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Issued March, 1954 Price Sixteen Shillings

SOME UPPER CRETACEOUS AND EOCENE
FRUITS FROM EGYPT

By M. E. J. CHANDLER

SYNOPSIS

The occurrence and preservation of fruits and seeds from the Dano-Montian and Eocene of
Egypt is described, and previous work summarized and in some instances revised.

These Egyptian fossils are clearly related to the London Clay Eocene plants on the one hand
and to the Recent flora of South-Eastern Asia on the other as shown by the occurrence of Nipa
in all three floras. The historical and geographical connexion of the three floras is considered,
due attention being paid to recent work on plant migration by Chaney and Axelrod. The
conclusion is reached that whereas all Tethyan (Indo-Malayan) types of flora may have their
remote origin in a uniform palaeotropical plant belt in Early or Middle Cretaceous times, some
secondary centre of colonization must be postulated for the Eocene tropical flora of Western
Europe where the original palaeotropical angiosperm flora must have been exterminated by
the Cenomanian transgression.

The obvious source of recolonization lay in South-Eastern Asia, because the great East-West
Tethyan ocean must have constituted a formidable barrier to mass migration from the African
continent. Moreover the marked Malayan relationship of the London Clay flora supports its
immediate Asiatic origin. Detailed systematic descriptions of the Egyptian fruits are given.

INTRODUCTION

A few fossil fruits recently discovered in Egypt were sent to the British Museum
for identification. They are of great importance, for hitherto the bulk of our
knowledge of the ancient flora of North Africa has been derived from the study of
wood (Krausel, 1939), whereas fruits and seeds were almost unknown. The majority
of the specimens came from the Dano-Montian Lower Esna Shales between latitudes
25° and 27° N. They were collected by two Egyptian geologists, Doctors M. Y.
Hassan and M. I. Youssef, who in appendices to this paper give details of the deposits
which yielded the fruits. (See also note on p. 187).

Dr. Hassan's finds were from the Kharga Oasis of the Western Desert, Dr.
Youssef's from the Kosseir area bordering the Red Sea.

Like the London Clay, with which comparison will be made in the following
pages, these plants occur in marine beds with a marine fauna, and must be presumed
to have been derived from the nearby land surface of the African continent a little
further south.

By a curious coincidence a fruit from the Lutetian (or possibly slightly younger
beds of Eocene age) of Egypt was also received for study at about the same time.
It was collected by members of the Anglo-Egyptian Oil Company, Messrs. Thiebaud
and Robson, at Wadi Rayan in the Western Desert, i.e., at about latitude 29° N. in
the Fayum. It is an extremely puzzling specimen, and some points about its anatomy
are rather obscure.

GEOL. II, 4. IZ

I5o UPPER CRETACEOUS AND EOCENE FRUITS FROM EGYPT

The preservation of these fruits and seeds is interesting and somewhat variable.
Some of them, e.g., Palaeowetherellia, are reproduced cell by cell in limonite in part
at least. Indeed they display the details of their structure almost as well as the
pyritized fruits of the London Clay. They have the merit of being more stable
and permanent in their preservation than pyritized specimens, but this advantage
carries with it the disadvantage that they do not so readily disintegrate and fall to
pieces along structural lines and natural planes of weakness as the London Clay
pyritized fossils so frequently do. Some of the best and most delicate evidence
from the London Clay fruits came from decaying specimens, not from those which
were artificially sectioned.

But some of the Egyptian fossils, e.g., Icacinaceae, have the appearance of being
purely internal casts, for although they were already chipped, no cell-structure was
exposed, or if visible was so indurated as to show little detail. Such specimens
are unlikely to provide more information if broken or sectioned, and it therefore
seems pointless to sacrifice them where clear and unmistakable evidence of internal
structure along natural planes of weakness is lacking. It is highly probable that
future collectors will in due course supplement the present meagre evidence from
such tantalizing casts.

Several of the new Dano-Montian specimens agree with one (¥.12985) which has
been in the collection at the British Museum (Natural History) since 1912. This
was described by Krausel (1939 : 106, pi. 2, figs. 9, 10 ; text-fig. 32) and was referred
by him to Diospyros schweinfurthi Heer. It came from the Lower Danian of Farafra
in the Western Desert at latitude 27° N. approximately. Heer's supposed Diospyros
and another specimen which he named Royena desertorum were originally described
by him in 1876 (see p. 168).

The examination of the new material, and more especially of a newly fractured
surface of ¥.12985, leaves no doubt that Heer's " Diospyros " and " Royena " both
belong to a single species which has no connection with the Ebenaceae. The species,
named hereafter Palaeowetherellia schweinfurthi (Heer), is obviously closely related
to Wetherellia variabilis Bowerbank from the London Clay, and to another species
W. dixoni (Carruthers) — to be redescribed — from the Bracklesham Beds of Selsey.
The genus is now referred tentatively to the family Euphorbiaceae.

Apart from the above, only six Tertiary angiosperm fruits from Egypt have
previously been recorded :

Palmacites rimosus Heer from the Danian of Kharga (Heer, 1876 : n, pi. i,
figs. 21, 22), an imperfect fruit of which no really distinctive characteristics are
shown or described. The anastomosing " fibres " in fig. 22 may be nothing more
than compression ridges such as are seen in places on the fruit of Icacinicarya
youssefi. Heer's determination must be regarded as very doubtful pending re-
examination of the specimen,

Securidaca tertiaria Engelhardt (1907 : 213, pi. 19, fig. 6) from the Eocene of
the Fayum. It would be unwise to comment upon this aceriform fossil without the
most careful scrutiny of the original material combined with a comprehensive study
of living fruits of this type in the various families in which they occur.

Nipadites sickenbergeri Bonnet from the Middle Eocene Nummulitic Limestone

UPPER CRETACEOUS AND EOCENE FRUITS FROM EGYPT

15*

(Marine Lower Mokattam of Gebel Giuchi, Mokattam), a species which the available
evidence identifies with Nipa burtini (Brongniart) (see pp. 159, 162).

Rubiaceocarpum markgrafi Krausel from the same beds which is clearly the same
Nipa represented by seeds with outer integument preserved (see pp. 161, 163).

Nymphaeopsis bachmanni Krausel from the Lower Oligocene of Cairo (Krausel,
I939 : 39» pl- 2» figs- 2-8 ; pi. 3, fig. 8 ; pi. 21, fig. 6 ; text-fig. 7) has features which
suggest that puzzling material has been wrongly interpreted. Further reference
is made to the matter on p. 183 when the structure of Thiebaudia rayaniensis is
discussed.

Teichosperma spadiciflorum Renner (1907 : 217 ; Krausel & Stromer, 1924 : 33,
pl. i, fig. 2 ; text-figs. 1-3) from the Lower Oligocene of the Fayum, referred tenta-
tively by Renner and with more certainty by Krausel & Stromer to Pandanaceae,
needs re-investigation especially as regards the number of locules and form of the
seed. If the seed is really curved as Krausel & Stromer's text-fig. 3 shows, the
family Myrtaceae should be explored. But before making confident statements
about this, it might be necessary to examine serial sections. Without further
evidence no really satisfactory determination can be made, although relationship
to Pandanaceae seems highly improbable.

It is not surprising that fruits from a remoter period of angiosperm history
than the Tertiary material hitherto studied are difficult to relate to living genera
or to place in Recent plant families. But in spite of the fact that it has not been
possible to identify all the specimens, the interest and significance of this tiny
flora are out of all proportion to its size.

The list of the newly found plants is given below.

Family

Nipaceae .
Anonaceae
Euphorbiaceae

Euphor-

biaceae ?
Icacinaceae

Flacourti-
aceae ?
Incertae sedis

Horizon and locality

Genus and species

Nipa burtini (Brongn.)
Anonaspermum aegypticum n. sp.
Lagenoidea trilocularis Reid &

Chandler
Lagenoidea bilocularis Reid &

Chandler
Palaeowetherellia schweinfurthi

(Heer)

Icacinicarya youssefi n. sp.
Icacinicarya sp. ?
Thiebaudia rayaniensis n. gen.

et sp.

Carpolithus hassani n. sp.
Carpolithus sp. (Icacinicarya ?)
Carpolithus sp.

Lower
Danian

Farafra

Dano-
Montian

Kosseir

Lutetian
(approx.)

Kharga and .
Kosseir
Kosseir

. Wadi Rayan
Kharga
Kosseir —

Fuller details of localities are given under the descriptions of the species in the systematic
part of this paper. No attempt has been made to determine a few small twigs,

152 UPPER CRETACEOUS AND EOCENE FRUITS FROM EGYPT

These discoveries are of outstanding importance for a number of reasons :

(1) As stated above, fossil fruits and seeds of any age are rare in Egypt. The
recent finds suggest that careful collecting, persisted in over a long period, would
reveal the presence of a considerable flora in that region.

(2) Very little is known about pre-Tertiary angiosperm fruits and seeds from
any area whatsoever. Knowledge derived from leaves or wood is usually comple-
mentary to that based on fruits and seeds rather than strictly comparable with it.
This was discussed by Reid & Chandler (1926 : 10-13) • Plant organs such as leaves
tend to be preserved in different deposits from fruits and seeds and to represent
different elements in the parent plant-formations. But here in Egypt are data
from pre-Tertiary Beds which are really comparable with the abundant Tertiary
records of fruiting organs elsewhere.

(3) The plants from Egypt are quite obviously related to the Tethyan type of
flora found in the London Clay. For even in so small a group as this under dis-
cussion similar or identical genera and even species occur. Note the presence of
Palaeow ether ellia (a distinct genus and species representing Wetherellia), of a Nipa,
of the two species of Lagenoidea actually found in the London Clay (so far as present
information can demonstrate), of an Anonaspermum, and of the family Icacinaceae.
Here is impressive evidence that the Malayan type of flora so characteristic of the
London Clay was already present in Africa towards the end of Cretaceous times.
It must be presumed to have flourished on the southern sea-board of the ancient
Tethys ocean.

(4) The presence in Egypt of a flora of strong tropical Malayan affinity, for such
must be the relationship of any flora at all comparable with that of the London
Clay, confirms the view supported by other lines of evidence that the climate of
Africa to the north of the equatorial belt must have been remarkably different
during the late Cretaceous and Eocene from what it is today. Nipa and its asso-
ciates demand not only warmth but a high degree of humidity (Reid & Chandler,
1933:74^5^.).

The discovery of this Tethyan flora in Egypt at the end of the Cretaceous must
inevitably raise the question, " What was its historical and geographical connection
with the biologically similar floras in the London Clay in Eocene times and in Indo-
Malaya at the present day?"

Did the Egyptian flora with the characteristic Nipa and its associates arise in
remote times in South-East Asia and spread thence into the African and European
continents as climatic conditions, combined with available migration routes, fav-
oured its expansion, or had it some other origin ? From what primitive source or
sources were these three related floras derived ?

Professor Kryshtofovich expressed the opinion that in the tropics of South-East
Asia the Malayan type of flora had remained " unmolested ever since its first descent
from its Cretaceous ancestors " (1929 : 310, 311).

Reid & Chandler (1933 : 82) stated that in their opinion the London Clay flora
had its origin in Malaya, whence it migrated northwards and westwards along the
shores of the Tethys ocean to Western Europe.

The late Professor Seward criticized the views of these three authors on the

UPPER CRETACEOUS AND EOCENE FRUITS FROM EGYPT 153

grounds that the early Tertiary flora of South-East Asia is but little known, and
that the evidence concerning it, if it exists at all, has not yet been published (1934 :
23), so that there may, for all we know, have been no Tethyan type of flora in Indo-
Malaya in the early Tertiary which could have served as a centre of dispersal.

He propounded two alternative suggestions as to the origin of the Tethyan flora
in Europe :

(1) That like the later Tertiary floras it may have reached Europe " not from a
South-Eastern home but from the North."

(2) That the London Clay flora may have been part of a widespread flora " which
in the course of the Tertiary period suffered progressive reduction and is now repre-
sented by enfeebled relicts in Indo-Malaya."

Before discussing these suggestions it may be of value to digress so as to take
account of recent important American contributions to plant history set forth by
Professor Chaney in two papers (1940, 1947). Chancy bases his views on successive
American fossil floras. These show that there was gradual displacement of temperate
to warm-temperate Cretaceous forests in low and middle latitudes in the United
States at the end of the Cretaceous period. Their place was taken by warm tem-
perate to sub-tropical vegetation (the Neotropical flora of Chaney) of Lower Eocene
age derived largely from the Antillean region on the East, and from Mexico and
Central America on the West. Evidently, he states, an early Tertiary northward
movement of plant populations as a whole occurred, so that the temperate forest
type of vegetation became concentrated in high latitudes, while tropical and sub-
tropical vegetation, now characteristic of forests near the Equator, spread into
middle latitudes, reaching 50° N. approximately on the western side of the continent,
and 37° N. on the eastern side.

Professor Chaney compares American plant history with that of the Old World,
and emphasizes that in approximately the same period a parallel story of plant
migration occurs there, modified to some extent by the extensive East-West barrier
of the Tethys ocean, which caused some differentiation of vegetation to the North
and South of it in Eocene and later times. He makes special mention of the tropical
Malayan London Clay flora at latitude 50° N. on the west side of the Eurasian conti-
nent whose modern equivalents (notably Nipa and mangrove) grow in the rain-
forest of Indo-Malaya. He also lists and shows on a map (1940 : 482, 483, text-fig,
i) numerous Eocene and a few Oligocene (palaeotropical) floras of the Old World
which have similar tropical relationship, while localities for the corresponding
(neotropical) floras in the New World are also plotted. He then recalls that since
the Oligocene a reverse movement has taken place in both hemispheres, the sub-
tropical and tropical forests gradually moving south again, giving way both in North
America and in Eurasia to that temperate vegetation which he calls the Arcto-
Tertiary flora.

In America the tropical vegetation has survived in the Antilles, the North of
South America, Central America and Mexico, with a few remains in South Florida.
In Eurasia it survives in south-east Asia.

Chaney perceives that such a wide-spread shifting of forest distribution must be
due to factors which were world-wide in operation, affecting whole plant populations

154

UPPER CRETACEOUS AND EOCENE FRUITS FROM EGYPT

(1947 : 141). If, as appears from the evidence available, he is right that migration
from the end of the Cretaceous onwards to the Oligocene was northwards, so that
plants from equatorial regions extended into middle latitudes, then the suggestion
made by Seward that the Eocene tropical floras, like the later ones, may have reached
Europe from the North does not fit into the picture and cannot be seriously enter-

chalazo

chalaza

position of

raphc

micropylc

hilum

incurving of Utta
to make a canal

radicle

chalaza

chalazo

TEXT-FIG, i. A-D Longitudinal sections through fruits showing placentation (diagram-
matic). A. Diospyros sp. Recent. B. Wetherellia variabilis Bowerbank. London
Clay ; Sheppey . c. Wetherellia dixoni (Carruthers) . Cuisian ? ; Selsey . D Palaeo-
wetherellia schweinfurth i (Heer) . Upper Cretaceous ; Egypt. E. Longitudinal section
through seed, Diospyros sp. Recent.

UPPER CRETACEOUS AND EOCENE FRUITS FROM EGYPT 155

tained. There is certainly no evidence in Europe of a Malayan flora in more
northerly stations than the London Basin.

Reid & Chandler (1933 : 62) were convinced by its past distribution and present
physiological requirements that the Nipa flora in the London Clay was living at the
most northerly limit of the climatic conditions (both of temperature and humidity)
which it could tolerate. Hence it is improbable that it could have come into being
in yet more northerly regions which would have been still more unfavourable in these
respects, regions which were, moreover, already occupied by the Arcto-Tertiary type
of vegetation suited to them. For these reasons the origin of the Malayan flora to the
North appears to be untenable.

Before considering Seward's other alternative suggestion, it may be helpful to
summarize briefly the facts (and deductions from them) which should be borne in
mind and correlated in any further consideration of these problems. They are as
follows :

(1) A late Cretaceous Tethyan flora in Egypt found at approximately 25° to 30° N.
probably derived from the African mainland at no great distance to the South.

(2) An early Eocene Tethyan flora in Western Europe extending North to latitude
50° approximately.

(3) The presence of similar tropical or sub-tropical floras in the Eocene and Oligo-
cene situated apparently along the former northern boundaries of the Tethys ocean,
as many of these floras yield Nipa.

(4) A living Tethyan type of flora restricted to South-East Asia (the Indo-Malayan
flora).

(5) A northward extension of the equatorial tropical forest belt between Cretaceous
and Oligocene times coinciding with the broadened latitudinal belt of increased
temperature and humidity.

With these facts and deductions in mind, we now turn to Seward's second alter-
native theory, viz., that the Tethyan flora was part of a widespread flora which
suffered progressive reduction, surviving in Indo-Malaya today. This is an un-
questionably true statement, for on actual fossil evidence, so far as it goes, the flora
of Indo-Malaya, or something closely akin to it, was once much more widely distri-
buted (Chaney, 1940 : 482-485, text-figs, i, 2 ; Edwards, 1936 : 28, text-fig. 9,
map showing the distribution of Nipa in relation to the approximate outlines of
the Tethys ocean) . But Seward's statement does not carry the matter far enough ;
it requires much more elucidation. Nor does it exclude the possibility of an imme-
diate (early Eocene and post-Cretaceous) Malayan origin for the palaeotropical
vegetation of Europe.

In support of the theory that the various Tethyan floras originated within a
uniform tropical belt of vegetation reference may be made to Axelrod (1952). He
believes tfrat in the Lower Cretaceous, " long distance migration may have been
more effective than at any later time," producing a more or less uniform type of
vegetation within given climatic belts.

During that period, he points out, the angiosperms were beginning to compete
successfully with the older waning Mesozoic floras which they were in process of
supplanting. He visualizes that by Middle Cretaceous times there must have been

156 UPPER CRETACEOUS AND EOCENE FRUITS FROM EGYPT

basic, more or less world-wide, angiosperm floras with the minimum of regional
differentiation, viz., a tropical to sub-tropical flora in lower and middle latitudes
more uniform in type throughout the world than has existed at any time since that
period, and a uniform temperate flora at higher latitudes.

It is not unreasonable to suggest that Axelrod's uniform tropical angiosperm flora
was the source from which the Recent tropical flora of South-East Asia and the
Cretaceous-Oligocene tropical flora of North Africa were derived by direct and un-
broken descent within these two regions. It may also have been the source of the
Eocene European tropical flora, not by direct descent in Western Europe but by a
more circuitous route. These suggestions are considered and explained in greater
detail below.

There is no reason to think that in Equatorial Asia there was ever a period during
the Tertiary when tropical conditions did not persist. Therefore it is not unreason-
able to postulate that the old palaeotropical flora probably survived over the
equatorial belt in that part of the world throughout Tertiary times, even after the
Oligocene, when it was driven south of its maximum Eocene extension. In that
equatorial belt it would have retained or gradually evolved a character of its own,
viz., that of the Indo-Malayan flora as we know it today. This is essentially the
opinion expressed by Kryshtofovich and quoted on p. 152.

In Africa the course of events may have been somewhat similar. Following upon
the contraction of the Tethys sea in Oligocene and later times and the resulting
climatic changes, the Tethyan flora must have been driven from the more northerly
part of its former Cretaceous and Eocene territory by the pressure of unfavourable
conditions. But it could probably have survived in the tropics of Central Africa.
Desiccation combined with the southern trend of plant migration in post-Oligocene
times would undoubtedly have prevented any later return to the more northerly
latitudes it had previously occupied. Deserts and the Tethyan sea (or its shrunken
remains) would probably have cut it off effectively from Asia. But within the
African equatorial belt it may have persisted, possibly giving rise (or giving place ?)
eventually to a distinctive African tropical vegetation as it evolved in isolation
from the Asiatic stream of life.

But what of the London Clay Eocene flora of Britain, and how was it related to
the uniform palaeotropical flora of Lower Cretaceous times ?

There can be no doubt that the Lower Cretaceous flora of this country must have
been eliminated by the great marine transgression of the Upper Cretaceous. This
transgression far overpassed the most northerly limits of tropical climate for which
there is any evidence, viz., about latitude 50° N. If the European Lower Cretaceous
palaeotropical flora persisted at all it must have been on islands, or on tracts of land
on the western borders of the present European continent, perhaps on land margins
now submerged beneath the sea. If Chaney is right about the world-wide character
of climatic changes, the flora could not, in all probability, have survived unless land
nearer the Equator was accessible for colonization during the late Cretaceous. For
in America, as we have seen, temperate and warm-temperate forests occupied low
and middle latitudes at this period, from which latitudes they were driven only at
the end of the Cretaceous by the northward march of Equatorial vegetation. It

UPPER CRETACEOUS AND EOCENE FRUITS FROM EGYPT 157

seems obvious that with the withdrawal of the Upper Cretaceous sea the re-exposed
land surface (now having a sub-tropical or tropical climate) would have been available
for recolonization by a flora of tropical type. This may have come either from
southern and western plant " reserves " as suggested above, or from the only two
alternative sources, viz., the African continent, or the south-east of Asia. In the
former we know that a Tethyan flora was present in the late Cretaceous. In the
latter there is no reason to doubt the existence of such a flora at that time as has
been pointed out above.

The theory of recolonization from the west offers no adequate explanation of the
marked Indo-Malayan aspect of the incoming Eocene flora. It is unlikely that
closely parallel development would have taken place independently in two such
widely separated areas. It is equally improbable that the source of recolonization
was the African continent. Had no serious physical barrier existed, the natural
solution of the problem would be to assume that by Eocene times the Cretaceous
Tethyan flora of Africa had migrated northwards and spread into Europe, as it
followed the extension in latitude of the hot and humid climatic belt favourable
to it.

But the existence of the great ocean barrier of the Tethys over perhaps 1,000
miles or more from South to North must have had a marked deterrent effect.

Axelrod (1952) discusses conditions affecting the migration of faunas and floras
in geological times. He maintains that plant distribution at any particular time is
" largely a reflection of the climates available for occupancy " (p. 177), i.e., climate
is the chief factor controlling plant migration. But whereas he considers that a
much higher degree of probability exists for plants than for animals in the matter
of over- water dispersal, he is also sure that " within any given climatic region distance
will finally impose a barrier to the migration of plants also." It is only fair to him
to state that in spite of this, he regarded the presence of palaeotropical Tertiary
vegetation on both sides of the Tethys as evidence that the sea in this particular case
(as well as in others) did not prevent the distribution of the flora on both sides of it
(1952 : 187). But this is a curious conclusion to have drawn, for he also explains
that " The water barrier that a continental flora can transgress within an epoch of
time (say, 10 to 15 million years) is not unlimited," while beyond 200 to 250 miles
the probability of colonization by a whole flora is low. Hence what he designates
the " waif assemblages " of ocean islands which lack balance, since whole classes
and orders expected are absent, while others are only poorly represented. " It
would appear," Axelrod writes, " that long-distance, over-water migration has not
been generally effective in populating widely separated continents." Even during
much longer periods of time, therefore, there may be good reason to doubt whether
the Tethyan flora in Africa could have migrated directly across the great ocean which
lay athwart its path to the North. Further, as Reid & Chandler (1933) pointed out,
the London Clay flora shows far less affinity with the flora of Africa than with that
of South-East Asia.

To the present writer, therefore, it seems highly improbable that wholesale migra-
tion took place across the Tethys.

There remains the third suggested source for the post-Cretaceous colonization of

158 UPPER CRETACEOUS AND EOCENE FRUITS FROM EGYPT

West Europe, namely the South-East Asiatic. On such evidence as is available
this appears to be by far the most probable source for these reasons :

(a) The Indo-Malayan region and Southern England both lay within the palaeo-
tropical climatic belt of Eocene times.

(b) The northern shores of the Tethys and the land adjoining were so situated
as to provide a possible migration route from South-East Asia to West Europe within
this greatly extended tropical belt of Eocene times which allowed the flora of equa-
torial regions to occupy many degrees of latitude North of its present limits.

(c) The London Clay flora is more closely linked with that of Indo-Malaya than
with that of any other existing flora.

The final answer to these speculations can only be provided by extensive re-
search on fossil floras from many widespread regions. At present the evidence is
insufficient to furnish incontrovertible proof, but the balance of probability seems
in favour of a remote origin of the Tethyan type of flora within a uniform palaeo-
tropical belt in the early Cretaceous, and of an immediate origin of the London
Clay flora by migration from Indo-Malaya between Cretaceous and early Eocene
times.

The subsequent history of the London Clay flora and the reasons why it survived
only in Indo-Malaya have been set forth by C. & E. M. Reid (1915), Reid & Chandler
(1933), Chaney (1940, 1947) and others. When during the Oligocene the connection
of the Tethys with the Indian Ocean was broken so that the former vast waterway
became a huge land-locked Mediterranean, while the great transcontinental mountain
barriers of Eurasia were being uplifted, progressive cooling of the climate in middle
latitudes occurred, so that more temperate plants from the North gradually supplan-
ted the former tropical ones of the Eocene. During this period final extermination
was the fate of the Tethyan flora in Europe. With the shrinking of the Tethys
a migration route to Asia no longer existed under favourable climatic conditions.
The direct route to the tropics in the South was blocked by the combined barriers
of East-West mountains and the East-West remains of the Tethys, as effectively
as Northward migration had formerly been blocked by the vaster Tethyan ocean of
late Cretaceous and Eocene times.

But in South-East Asia, as already stated, survival of the palaeotropical flora
and its descendants was possible, for there is no reason to think that equatorial
latitudes in that continent have ever passed through a phase when it could not
have retained its tropical flora. Even when the more northerly parts of the palaeo-
tropical belt became cooler, so that the tropical flora could no longer live in them,
migration to more southerly regions was possible, thanks to the absence of East-
West barriers and to the existence of North-South valleys and coastal plains.

Meanwhile it is likely that the Tethyan flora of Egypt, when more fully known,
will reveal a far stronger African element than is found in the London Clay, cut off
as this latter appears to have been from direct communication with the African
continent throughout its history.

The affinities of unknown plants in the Egyptian fossil flora should therefore be
sought among tropical and sub-tropical African genera of the present day as well
as among those of South East Asia.

UPPER CRETACEOUS AND EOCENE FRUITS FROM EGYPT 159

I wish to express my warmest thanks to Mr. W. N. Edwards for helpful suggestions
and criticisms, and to Mr. F. M. Wonnacott of the Geological Department for much
help in the preparation of the manuscript for the press.

SYSTEMATIC DESCRIPTIONS

Note. — The specimens, with two exceptions, are from the Dano-Montian Lower
Esna Shales of Egypt.

Those from Kosseir Area, Red Sea, were collected by Dr. M. I. Youssef, those
from Kharga Oasis by Dr. M. Y. Hassan, to both of whom I am indebted for the
opportunity to study these most interesting plants and for the generous gift to the
British Museum of the figured specimens.

The other two horizons and localities represented are the Lower Danian of Farafra
(Palaeowetherellia) , and the Eocene beds of Wadi Rayan (Thiebaudia rayaniensis
gen. et sp. nov.).

MONOCOTYLEDONES

Family NIPACEAE
Genus NIP A Thunberg 1782

Nipa burtini (Brongniart)
(PI. 10, figs. 1-5 ; pi. n, figs. 7-9)

1904. Nipadites sickenbergeri Bonnet, p. 499, figs, on pp. 500, 501.

1924. Nipadites sickenbergeri Bonnet : Krausel, p. 36.

1939. Nipadites sickenbergeri Bonnet : Krausel, p. 22, pi. I, figs. 1-18, text-fig. I.

!939- Rubiaceocarpum markgrafi Krausel, p. 108, pi. i, figs. 19-24.

For full synonymy see Reid & Chandler (1933 : 118).

LOCALITIES AND HORIZONS. North slopes of Mokattam, near Cairo ; Middle
Eocene. Gebel Atshan, Kosseir Area, Red Sea ; Dano-Montian Lower Esna Shales.

DESCRIPTION AND REMARKS. The discovery of a small Nipa. fruit in the Dano-
Montian of Kosseir and the re-investigation of fruits named Rubiaceocarpum mark-
grafi Krausel from the Middle Eocene of Mokattam made it necessary to re-study
Nipadites sickenbergeri Bonnet from the same beds as Rubiaceocarpum.

As a result it can be stated with confidence that Rubiaceocarpum is a Nipa seed
with outer integument preserved in part at least, and that it, and the seeds named
Nipadites sickenbergeri, cannot be separated from Nipa burtini (Brongn.) in the
present state of our knowledge.

This conclusion was forced by the evidence upon the writer who had started with
the conviction that N. sickenbergeri was distinguished beyond question from Nipa
burtini by its ribbed and furrowed seed. Moreover the opinion expressed by Reid
& Chandler (1933 : 122) that the fossils should be referred to Nipa rather than to
Nipadites is now re-affirmed.

160 UPPER CRETACEOUS AND EOCENE FRUITS FROM EGYPT

In order the better to understand the fossils a further examination of the living
Nipa fruticans was carried out. The following details may now be added to those
recorded by Reid & Chandler (1933 : 122).

The hard compact endocarp when freed from the adherent pericarp (or sarcocarp)
is more or less smooth but shows on its outer surface, in the parenchyma which
covers it, impressions of stout longitudinal fibres. There is no conspicuous fluting
or ribbing of this surface, but a tangled mass of finer fibres can be seen where the
parenchyma is rubbed off. When abrasion is carried a stage further stout transverse
fibres are exposed. These were observed in 1933. The locule shows conspicuous
flattened transverse or oblique fibres and stout longitudinal ones overlying them
which readily break away.

When an endocarp is cut transversely, a nut-like seed may fall free if the material
is in a suitable state of preservation. This bears upon its surface conspicuous flutings
due to narrow deep furrows in a parenchymatous layer. In the furrows there are
remains of fibres which probably belong to the endocarp. They have usually been
torn away, leaving empty furrows only on the surface of the seed. Some of the
fibres branch or unite so that the furrows and intervening ridges are not aways
absolutely straight and longitudinal, although this is their general alignment. On
the flat or slightly convex broad ridges between the furrows can be seen impressions
of the transverse or oblique flattened fibres which belong to the locule wall described
above (PI. n, fig. 10).

In Reid & Chandler's description this coat of the testa was regarded as part of
the endocarp because endocarp and testa are often fused. On the detached seed
there is a deep narrow parallel-sided furrow (PL n, fig. 12). This is formed by an
incomplete longitudinal ridge or septum which projects into the locule from the
whole length of the carpel wall. The large sub-circular or circular basal aperture
giving access to a cavity in the endosperm is a conspicuous feature also, and is
associated with the embryo. The outer pulpy integument of the testa is fairly
easily scraped away, exposing a close-textured rather brittle, gummy or resinous
inner integument with a smooth transversely striate outer surface of fine cells.
Embedded in the surface of this coat are a few flat fibres which branch and anas-
tomose to form a coarse network. The fibres lie obliquely, transversely, or more
infrequently longitudinally (PI. 10, fig. 6). On the inner side of the brittle inner
integument is a raised network due to very fine fibres so arranged as to form fine
transversely elongate meshes.

The inner integument follows fairly closely the contours of the endosperm, but is
separated from it by a thin soft film of fine cells. When the endosperm has rotted,
as may happen in specimens which have drifted in the sea for some time, the empty
integuments of the seed may still remain attached firmly to the endocarp as it lies
within the drupe, but the film of cells just described has usually gone so that only
the network of fibres with transverse meshes covers the exposed surface. It is the
internal cast of this coat with its fibres which shows the ornamentation described
on seed-casts of Nipa burtini from the London Clay (Reid & Chandler, 1933, pi. 2,
fig. 6).

No raphe or chalaza scar is visible on the interior of the inner integument in

UPPER CRETACEOUS AND EOCENE FRUITS FROM EGYPT 161

Recent fruits, nor is their existence indicated by any change in the alignment of the
fine fibrous network so far as I have been able to observe. When all the integuments
are removed so that the actual endosperm is uncovered, a series of even finer furrows
and ridges, like minute ruminations, are laid bare. At the apex and over the sides
of the endosperm their orientation is clearly transverse (PI. 10, fig. 6 ; PI. n, fig. n).
At the base they are coarser and the alignment more irregular, giving rise to small
low rounded lobes of endosperm. Whatever the orientation, the small areas which
they produce on the surface are more or less convex, especially when slightly weath-
ered or abraded. The endosperm sometimes displays a few broad shallow longi-
tudinal furrows or sinuosities at the lower end which gradually die out above (PI. II,
fig. 12). They resemble the furrows sometimes seen in Nipa burtini and Nipadites
sickenbergeri. On the exterior of the complete seed they are barely discernible,
being obscured by the spongy outer integument. Nor are they usually apparent
on the inner surface of the empty testa.

The astute observer Hooker in an editorial note (Le Maout & Decaisne, 1876 :
822) comments that the seeds are erect and anatropous. But for this observation
I should not have suspected their anatropous character, for there is no more indi-
cation of a raphe on the outer surface of the testa than there is on the inner surface.
Nevertheless examination of a detached endosperm-mass supports Hooker's state-
ment, or at least gives evidence of comparable structures. After removal of the
integuments a flat band of fibres was seen lying in one of the shallow longitudinal
furrows of the surface. It extends upwards from the base almost throughout the
length, follows a straight course, and is parallel-sided, quite different in character
from the somewhat sinuous flat fibres in other furrows. That this is a definite
structural feature, probably the raphe, is confirmed by the differentiation of a band
of endosperm immediately beneath it. In this position there are small irregular
rounded " ruminations," not transversely elongate ones as elsewhere, and some
indication of longitudinal alignment is given by a tendency for longitudinal cracks
to form (PI. n, fig. n). Somewhere in this band the fibres must enter the endo-
sperm and pass to the chalaza. But the point of entry is so inconspicuous that it is
hidden by the irregularities of the " ruminate " surface. Nevertheless a correspond-
ing structure shows clearly in some fossil material, as will be described later.

From the details given above it is fairly clear that the appearance of a fossil
Nipa must vary considerably with the coat which happens to have been preserved
or exposed. This depends partly on the mode of preservation of the fossil. Thus
an internal seed-cast may show the structure of the endosperm as in the Egyptian
Nipa, or it may merely be a cast of the inner integument formed after the endosperm
had decayed as in many London Clay seeds.

Any seed having the ridged outer integument preserved would have the dis-
tinctive appearance of RuUaceocarpum, further details of which are given later.
In a broken fruit some layer or layers of the endocarp may show, as in many fruits
from the London Clay, and in a small Dano-Montian fruit from Egypt. In the
London Clay fruits wear and tear has often exposed the inner layers of endocarp
with flat transverse fibres described in 1933. By far the most common specimens
at Sheppey and Herne Bay are fruits (perfect or imperfect), showing stout fibres

i6z UPPER CRETACEOUS AND EOCENE FRUITS FROM EGYPT

embedded in the spongy tissue of the sarcocarp. Rarely is the epicarp preserved,
and then only in fruits freshly removed from the matrix. It quickly cracks and
falls to pieces on exposure to air. .

It seems scarcely necessary to repeat detailed descriptions of the fossil fruits and
seeds which have already appeared in published work (Reid & Chandler, 1933 : 119 ;
Krausel, 1939 : 22 ; Seward & Arber, 1903, etc.). Other references will be found
in the synonyms listed by Reid & Chandler (1933 : 118).

The question inevitably arises once more, " How many fossil species can be dis-
tinguished in the deposits with which we are now concerned ?"

It must be remembered that whereas the London Clay fruits and seeds are rela-
tively uncrushed, the Egyptian ones have often undergone an excessive amount of
deformation by crushing and there appear to be no valid and consistent grounds for
separating Nipa burtini and Nipadites sickenbergeri as distinct species. The longi-
tudinal ridging and furrowing of the seed-casts, or its absence, although it was
supposed by Bonnet to be of specific value, cannot really be used in diagnosis, for
although smooth seed-casts are common in the London Clay, furrowed ones may also
occur. Moreover both kinds are found in the Egyptian Eocene (Bowerbank, 1840,
pi. 4, fig. i; pi. 5, fig. i ; Krausel, 1939, pi. i, figs. 11-13).

Again in the Egyptian casts the furrows may be few and slight (V. 13695) or fairly
conspicuous, or in some cases, overemphasized by crushing.

The other chief difference which occurs in some cases between fossils from the
two localities lies in the size. But differences in size are no more satisfactory than
furrowing, as a reason for specific separation. It is true that the majority of Nipa
burtini fruits and seeds from the London Basin are smaller than the majority of the
seeds from Egypt, and that they vary greatly in Appearance. Yet large nuts do
occur in the London Clay (Bowerbank, 1840, pi. 5, fig. i), while very large fruits
and seeds occur in the Belgian strata (Seward & Arber, 1903). On the other hand
the small fruits from the London Clay are indistinguishable from a Dano-Montian
fruit from Kosseir (PL 10, figs, i, 2). Perhaps the predominance of small forms in
one area and large in another may be due to the sorting effects of specific gravity
rather than to systematic differences. The associated fruits of other families at
Sheppey and Herne Bay are on the whole fairly small. .

The following table shows minimum and maximum sizes of fossil Nipas from
previously published records. It must be remembered, however, that a far larger
number of specimens are known from the London Clay than from the other deposits
and they include many immature or abortive fruits.

London Clay Belgian Eocene Egyptian Eocene

Fruits :

Length, i to 18 cm. . 7-5 to 21 cm. Not known

Breadth, i -3 to 12 cm. . 3 to 15 cm. . „

Seeds :

Length, 2 to 8-9 cm. . 9 to 10 cm. (commonly 8-5 . 7-5 to n cm. (commonly 8-5

to 9 cm.) or 9 cm.)

Breadth, 1-3 to 7 cm. . 9 cm. (commonly about 7 . 3-4 to 12-3 cm. (commonly

cm.) about 5 to 7 cm.)

UPPER CRETACEOUS AND EOCENE FRUITS FROM EGYPT 163

Thus whether furrowing or size be considered, there appear to be no sharp lines
of demarcation between species from the localities with which we are concerned.

If furrowing and size cannot be used as a basis for separating species — and it is
difficult to see how they can when every gradation may occur in these characters —
there are no other grounds obvious on which the diagnosis of more than one species
can be based.

Certainly the differences between the Nipas from the two areas under review
are no greater than the differences among individual specimens in the London Clay
itself (cf. Bowerbank, 1840, pi. 4, fig. i ; pi. 5, fig. i). Yet there is no reason whatso-
ever for creating more than one species in this deposit.

A few additional observations on the Egyptian nuts may not be out of place
here. Krausel's figured specimens are largely seed-casts and might be expected
to show endosperm structure. Unfortunately the small scale of the figures does not
display such fine details except in his pi. i, fig. 8, where transversely aligned endosperm
cells are clearly seen around the basal scar. Endosperm structure is also clearly
visible in two casts from Mokattam in the British Museum (V. 13240 and ¥.13695 ;
PI. n, figs. 7, 8).

The variable and considerable amount of compression of the seeds must be stressed,
for some are almost reduced to thick concavo-convex lenticles (¥.13240). Inevitably
this crushing increases the diameter. ¥.13239 is obliquely distorted, the basal
aperture having been forced into a basi-lateral position (PI. 10, fig. 3).

As in the living, so also in the fossils, endosperm may show features not dis-
cernible on the interior of the testa represented by internal casts. ¥.13240, in
addition to the typical transverse pattern of endosperm ridges (PL n, fig. 8), shows
on its flatter surface a band of elongate, longitudinally aligned endosperm cells
which by comparison with the living have structural significance, probably indicating
the position of the raphe. Towards, but well below, the apex a radial arrangement
of the endosperm on this band suggests the point of entry into the endosperm of the
fibres from the raphe (PI. n, fig. 7). On the more convex surface of this specimen,
the coarse network of fibres seen in the living between the two integuments of the
testa are faintly impressed (PL 10, fig. 5).

¥.13239 has a mosaic pattern all over the surface. This may be the impression
of a cracked testa which has peeled. But the important feature of this specimen is
that sub-apically the cast shows a deep, small, funnel-shaped opening towards
which fibre-impressions converge, and into which they pass (PL 10, fig. 4). Here
the entry of the raphe fibres is clearly indicated. ¥.13695 shows a similar but more
obscurely preserved radiating structure.

Thus these specimens illustrate a well-established fact, that owing to the macera-
tion that fossils have frequently undergone, they may display structures which are
more difficult to demonstrate in Recent material.

We must now return to the specimens named Rubiaceocarpum markgrafi Krausel.
As already stated on p. 159 these appear to be seeds (abnormally short and broad)
with outer integument preserved. They have undergone considerable lateral com-
pression.

Their form can be matched among the Nipa seed-casts illustrated by Krausel

GEOL. II, 4. I2

164 UPPER CRETACEOUS AND EOCENE FRUITS FROM EGYPT

(1939, pi. i, cf. figs. 8, 21 ; 18, 24 ; 16, 24 ; 17, 20). I have been able to examine
the specimen illustrated in Krausel's fig. 24, which shows the base of the seed with
its large aperture. A very interesting feature is a broad flat fibre band which half
encircles the seed in the plane of symmetry (PI. n, fig. g/). This resembles the
raphe fibre described above in Nipa fruticans, although seed and fibre are on a
larger scale. The fact that it is visible at the surface of the specimen must mean
that some measure of abrasion had removed part of the spongy parenchyma. The
superficial impressions of the transverse fibres of the endocarp have also been des-
troyed at the same time, traces only of them being visible in a few of the deeper
furrows of the outer integument.

We may ask what relationship does this ribbed entity bear to its living analogue?

It appears to be an internal cast of a hollow mould left by a seed which subse-
quently decayed. The filling of the hollow with limestone resulted in the characters
impressed on the mould being reproduced on a cast, much as sealing-wax poured
into the hollow mould of any fossil will reproduce its solid form. An actual replace-
ment of the seed itself cell by cell would have given a far sharper representation of
it, and in places at least would have provided evidence of cell-structure.

The ribbed seed shown by Krausel (1939, pi. i, figs. 20, 21) perhaps retained even
more of the outer integument, for there are suggestions of transverse fibre impres-
sions at the base of fig. 21, and the raphe band appears to be only partly uncovered.

The longitudinal striation on the ridges in Krausel's pi. i, fig. i, may indicate
that part of the outer integument of the testa is still present on this cast, which
he referred to Nipadites. But suggestions such as this can only be verified by
examination of the specimens themselves.

Reid & Chandler (1933 : 122) gave reasons for referring the London Clay fossils
to Nipa rather than to Nipadites. They appear still to hold good. The most out-
standing difference between living and fossil is the presence of the short longitudinal
septum, which by partial subdivision of the locule in N. fruticans produces the
only constant deep and conspicuous longitudinal furrow on the seed. This septum
and furrow are absent in all the fossils. Nevertheless so closely identical with the
living is the structure of the fossils in all other respects (even in smallest details)
that the presence or absence of the septum still appears to the writer to have specific
rather than generic value. Probably however, the real meaning of this character
can only be decided by a detailed study of the developing ovary. It may, or may
not, have a connection with the suppression of two out of the three original locules
or ovules. In the meanwhile it is suggested that the name Nipa be used for the
fossils.

UPPER CRETACEOUS AND EOCENE FRUITS FROM EGYPT 165

DICOTYLEDONES

Family ANONACEAE

Genus ANONASPERMUM Ball emend Reid & Chandler, 1933

Anonaspermum aegypticum n. sp.

(PL n, figs. 13-15)

DIAGNOSIS. Seed-cast oval or oboval in outline, much compressed with slight
median depression ; rumination ridges close and narrow, extending from the depres-
sion to the margins, diverging from the depression at the distal end, occasionally
forking near the margin, with a few short intermediate ridges. Four-partite in
transverse section. Thickness only about half the greatest diameter in the plane
of symmetry (15-5 mm.).

HOLOTYPE. A seed-cast, distal end, with testa almost entirely abraded (PL n,
figs. 13-15). Brit. Mus. (N.H.), No. ¥.31106.

LOCALITY AND HORIZON. Gebel Atshan, Kosseir Area, Red Sea ; Dano-Montian
Lower Esna Shales.

DESCRIPTION. The unmistakable internal cast of a seed of Anonaceae showing
typical ruminate albumen and marginal encircling band of raphe and chalaza
(PL n, figs. 13, 14). The hilar half of the seed is missing. The surviving fragment
represents the distal end, which appears to have been severed from the proximal
only a short distance above the middle of the specimen. The cast was originally
oval or oboval in outline, bisymmetric, somewhat compressed at right angles to
the plane of symmetry, anatropous with encircling raphe in the plane of
symmetry. The raphe begins to broaden on one side where it merges into the
linear chalaza (PL n, figs. 13, 14 ch.}. It completely fills the slight marginal
groove at the edge of the albumen. Albumen having a median depression on each
broad surface, ruminations forming narrow close ridges from the margin of the seed
to the median depression. The ridges diverge from the end of the depression to
the apex or distal end of the seed. The inner ends of opposite ridges do not unite
at this point to form a series of curves as in Anonaspermum commune Reid & Chandler
(1933 : 184, pi. 5, figs. 14-17). Short ridges of varying length sometimes arise at
the margin and are interposed between the longer ones. Occasionally one of the
longer ridges may divide into two towards the margin of the seed. The transverse
section exposed on the fractured surface shows a four-partite arrangement of the
albumen (PL n, fig. 15). In the slight depressions in the middle of the broad
surfaces part of the fibrous layer of testa is preserved (PL n, fig. 13). On the four-
partite sectioned surface oblong cells arranged in transverse rows at right angles to
the plane of symmetry of the seed can be seen. The rows are about 0-028 mm. broad.
The cells are partially obscured by sinuous fine striations lying parallel in a general
way with the plane of symmetry. Parts of the ruminations as seen on the fractured

166 UPPER CRETACEOUS AND EOCENE FRUITS FROM EGYPT

surface of the seed are obscured by a thin brown mineral film which bears the
impression of a thin but rough coat, the cells of which lie at right angles to the plane
of symmetry. It represents the filmy inner integument which penetrated between
the plates of albumen as in living seeds.

Actual length of seed preserved, 10 mm. ; estimated length of complete seed,
about 22 mm. Maximum diameter actually preserved in plane of symmetry,
14 mm. ; probable measurement in the complete seed, about 15 to 15-5 mm. Dia-
meter at right angles to plane of symmetry, 7 (at the centre) to 7-5 mm. (at each side).

REMARKS. The seed was clearly larger than any described from the London
Clay, about twice the size of the largest there recorded. It was also relatively
thinner at right angles to the plane of symmetry than any London Clay species
with albumen forming superficial ridges. Similar divergence of the ruminations at
the distal end is seen in A. rugosum and A. pulchrum Reid & Chandler (1933 : 186,
187, pi. 5, figs. 21-27), but not in A. commune and A. rotundatum Reid & Chandler
(1933 : 184-187, pi. 5, figs. 14-20).

Although imperfect, this solitary specimen appears to be sufficiently distinctive
to merit a specific name. It is described as Anonaspermum aegypticum. The family
Anonaceae is .almost exclusively tropical today, occurring in both hemispheres. It
is recorded from both hemispheres in Eocene times also.

Family EUPHORBIACEAE
Genus LAGENOIDEA Reid & Chandler emend.

Since the first description of this genus was published (Reid & Chandler, 1933 :
493-497) a few small but important new facts have come to light which make it
necessary to correct the diagnoses and descriptions then given. The information
was derived from additional London Clay material of Lagenoidea trilocularis. The
diagnosis should now read :

Fruit a superior, syncarpous, loculicidal and septicidal capsule, two to four-
loculed, locules single-seeded. Pericarp thick, formed of radially aligned cells,
seeds pendulous, radially compressed, anatropous, raphe ventral, chalaza basi-
ventral.

TYPE SPECIES. L. trilocularis Reid & Chandler.

The recent discovery that the seeds are pendulous with ventral raphe now makes
it possible to refer the genus Lagenoidea to the family Euphorbiaceae. Formerly
such a relationship was considered impossible, for the seeds were then believed to be
erect and orthotropous owing to the fact that the raphe is slender and difficult to
detect when it is preserved at all.

The shape of the fruit and the combined loculicidal and septicidal dehiscence is
typical of the Euphorbiaceae. This was recognized by Ettingshausen when he
labelled one specimen (¥.23129) Euphorbiophyllum eocenicum (see Reid & Chandler,
1933 : 495, pi. 29, fig. 6). This specimen was never described or figured as such,
hence the inappropriate name Euphorbiophyllum need not be retained for these

UPPER CRETACEOUS AND EOCENE FRUITS FROM EGYPT 167

fruits. It has not yet been possible to connect them with a living genus, but the
relationship must be sought among those sections of the Euphorbiaceae with single-
seeded locules having locule-linings formed of complicated interlocking cells. Attenu-
ated obovate seeds are rare in the family but obovate seeds may occur, e.g., in
Chaetocarpus where the seeds may remain attached to the inner angle of the carpel
wall when the external valves have fallen away.

Lagenoidea trilocularis Reid & Chandler

(PI. 12, figS. l6-20)
1933. Lagenoidea trilocularis Reid & Chandler, p. 493, pi. 29, figs. 1-18.

The revised diagnosis of this species based on London Clay material as stated
above should read :

Fruit : Sub-globular, three- (rarely four-) loculed ; capsule splitting loculicidally
and septicidally into six (rarely eight) segments. Length, 7 to 14 mm. ; diameter,
8 to 23 mm. ; length of locule-cast (= seed), 3-5 to 4-5 mm. ; greatest diameter,
2 mm. ; least diameter, i mm.

LOCALITY AND HORIZON. Gebel Atshan and Gebel Durvi, Kosseir Area, Red
Sea ; Dano-Montian Lower Esna Shales.

MATERIAL. Two fruits, one with hollow interior. Brit. Mus. (N.H.), Nos.
¥.31107-08.

DESCRIPTION AND REMARKS. Two fruits, neither of which show internal structure.
One (PI. 12, figs. 16-18) shows a thin wrinkled epicarp partly hiding the surface of
the capsule, but the six segments can be clearly distinguished. A sub-circular
basal scar indicates the former extent of the calyx (PI. 12, fig. 17). Length of fruit,
10 mm. ; transverse diameter, 9 by n mm.

The second fruit (PI. 12, figs. 19, 20) also shows the six valves and basal scar,
but its surface is somewhat encrusted and is therefore rather obscure. The interior
is hollow, while there is a hole at the attachment through which the seeds may be
presumed to have escaped. Length of fruit, n mm. ; diameter, 14 by 12-5 mm.

Lagenoidea bilocularis Reid & Chandler
(PL 12, figs. 21-23)

1933. Lagenoidea bilocularis Reid & Chandler, p. 496, pi. 29, figs. 19-27.

LOCALITY AND HORIZON. Gebel Atshan, Kosseir Area, Red Sea. Dano-Montian
Lower Esna Shales.

MATERIAL. Two typical fruits. Brit. Mus. (N.H.), No. ¥.31109.

DESCRIPTION. Two sub-ovoid somewhat laterally compressed fruits showing
six external longitudinal grooves delimiting the valves and septum. The narrow
median opposite segments between the pairs of lateral segments mark the edges of

168 UPPER CRETACEOUS AND EOCENE FRUITS FROM EGYPT

the septum. The pairs of end segments are bounded by planes of loculicidal and
septicidal splitting. As in the London Clay fruits of this species the pairs of valves
may be presumed to overlie the two locules.

The stylar scar is seen at the narrow end in both fruits and the attachment scar
is preserved at the broad end in the smaller specimen. Unfortunately the internal
structure is not exposed. There can be no doubt however that the fruits are identical
with the London Clay species Lagenoidea bilocularis Reid & Chandler.

Length of the two endocarps, n and 9-5 mm. ; greatest diameters, n and 9 mm. ;
least diameters, 6 and 5-5 mm. respectively.

Family EUPHORBIACEAE ?
Genus PALAEOWETHERELLIA nov.

DIAGNOSIS. Fruit sub-globular, syncarpous, with more than five carpels ; dehis-
cence loculicidal and septicidal, locules radially arranged, tangentially compressed ;
placentation axile ; seeds solitary in each locule, occupying part only of the plane
of the locule ; pendulous by long arched funicles which arise some distance below
the apex of the fruit. Pericarp parenchymatous. Seeds slightly inflated, obovate
in outline, beaked at the micropyle, anatropous with ventral raphe. Testa one
cell thick, formed of equiaxial cells.

TYPE SPECIES. P. schweinfurthi (Heer).

Palaeowetherellia schweinfurthi (Heer) Chandler

(PL 12, figs. 24-30 ; PL 13, figs. 31-38 ; PL 14, figs. 39-47 ; PL 15, fig. 48 ;

Text-fig, i)

1876. Diospyros schweinfurthi Heer, p. 6, pi. i, figs. i-io.

1876. Royena desertorum Heer, p. 10, pi. i, figs. 11-16.

1889. Diospyros schweinfurthi Heer : Schenk, p. 745, text-fig. 384 7~13.

1889. Royena desertorum Heer : Schenk, p. 745, text-fig. 38414~18.

!939- Diospyros schweinfurthi Heer : Krausel, p. 106, pi. 2, figs. 9, 10 ; text-fig. 32.

1939. Royena desertorum Heer : Krausel, p. 106.

DIAGNOSIS. Fruit shorter than broad, 6 to 9, 10 or even 12-loculed. Funicles
arising from the axis at about one-third to one-half the length of the fruit from
the apex, passing between the closely opposed surfaces of the locule above the seed.
Seed so obliquely placed as to lie almost transversely with the hilum upwards to-
wards the circumference of the fruit, and the ventral margin parallel with the arched
funicle. Fine parenchyma which forms the bulk of the carpel wall enclosing numer-
ous patches of coarser cells very liable to decay associated with a system of fibres
near the surface of the endocarp. Diameter of endocarp, 17 to 22 (or ? 24) mm. ;
length about 7-5 to 10 mm. Maximum length of fruit with exocarp preserved, 14
mm.

UPPER CRETACEOUS AND EOCENE FRUITS FROM EGYPT 169

NEOTYPE. A nine-loculed fruit (PL 13, figs. 31-37). Figured Krausel, 1939,
pi. 2, figs. 9, 10 ; text-fig. 32. Brit. Mus. (N.H.), No. ¥.12985.

LOCALITIES AND HORIZONS. Farafra, Egypt ; Lower Danian (Upper Cretaceous).
Gebel Tarawan and Gebel Um-el-Ghanaim, Kharga Oasis, Egypt ; Dano-Montian
Lower Esna Shales. Gebel Atshan, Gebel Durvi and Abu Tundub, Kosseir Area,
Red Sea ; Dano-Montian Lower Esna Shales.

DESCRIPTION. Fruit : Having an exocarp (leathery ? but usually abraded)
about 1-6 mm. thick, its surface more or less smooth but the details of its structure
obscured by mineral incrustation (PI. 12, fig. 27 ; PI. 14, figs. 42, 43). It encloses
a syncarpous multilocular endocarp, circular or sub-circular in outline, sometimes
slightly angled over the locules, depressed dorsiventrally, usually somewhat flattened
at the apex, either slightly pointed below or somewhat excavated (PI. 12, figs.
24-26 ; PL 13, figs. 31, 32).

The style base may be marked by a small inconspicuous scarcely prominent cir-
cular scar (PL 13, fig. 31), but this is not usually apparent. External surface with
radial ridges or sutures which correspond with the locules. Alternating with them
are less conspicuous ones which overlie the septa ; they are not invariably seen, and
are most marked in much abraded specimens or in those which have started to split
septicidally (PL 12, figs. 25, 29, 30). Dehiscence conspicuously loculicidal, less
obviously septicidal. Possibly the septicidal planes of weakness are cemented by
infiltration of mineral substance as they are normally so inconspicuous. Carpels
6 to about 12, radially arranged about the axis of the fruit (PL 12, figs. 24-26 ;
PL 13, figs. 31, 32), unequally developed in some specimens so that complete fruits
may show perfect or imperfect radial symmetry. ¥.12985, for example, shows
unequal development at opposite ends of one diameter, two of the smaller carpels
are evidently abortive, but their existence is clearly revealed by a longitudinal
fracture of the fruit (PL 13, figs. 33-36). One of Heer's transversely sectioned
specimens (1876, pi. i, fig. 9) in which he only recognized 8 locules clearly shows 9
in the figure, 2 being abortive or ill-developed. Up to the present the least number
of carpels seen is six (PL 12, figs. 24, 25). Fibres of the fruit axis fused with the
surrounding parenchyma not occupying a distinct central canal. Locules extend
from the axis to the periphery of the fruit, but are tangentially compressed and so
completely flattened around their edges that the two flat surfaces are contiguous.
The seeds are neither as long nor as broad as the locules. It is only where the seed
actually lies that there is a slightly inflated cavity equal to the thickness of the seed
(PL 13, figs. 32-36 ; PL 15, fig. 48). Placentation axile. Seeds solitary, suspended
by long arched funicles which spring from the axis at about one-third or one-half
of the length of the fruit from the apex. From the point of emergence of the funicle
from the axis (PL 13, figs. 33-37 ; Text-fig. ID) to the point where it passes into the
seed at the hilum it lies within that part of the locule where the opposed walls are
contiguous. The longer axes of the seeds lie at an angle of 60° or thereabouts to the
axis of the fruit, hence the, position of the seeds is oblique or almost transverse in
the fruits. As this is a consistent feature it must be original. Owing to this
peculiarity, the dorsal surface of the seed lies towards the base of the fruit, the hilar
end is directed outwards and upwards, the distal end lies close to the lower end

170 UPPER CRETACEOUS AND EOCENE FRUITS FROM EGYPT

of the fruit axis, and the ventral margin is uppermost, lying parallel with the long
arched oblique funicle.

Pericarp of three layers : (i) a leathery (?) exocarp of fine parenchyma whose
surface is obscure, but in section it can be seen in one place to be formed of rounded
more or less equiaxial cells about 0-028 mm. in diameter. (2) A compact parenchy-
matous coat which constitutes the main thickness of the endocarp. It is greatly
thickened in the part of the fruit between the flattened margins of the locules.
The outermost part of the endocarp is formed of close fine parenchyma which is
rather readily abraded. Superficially, when the coat is weathered, the cells can be
seen to be arranged in radiating groups. Possibly there is a fibre at the centre of
each group. Differential weathering of this coat produces a series of rugosities
having the effect of a coarse network, the centre of each radial group being sunk
below the general level of the surface. Beneath the layers of radially grouped cells
comes the main thickness of the endocarp. It is formed of equiaxial cells about
0-02 mm. in diameter. Within it are numberous patches of coarse angular paren-
chyma, with cells about 0-05 to o-i mm. in diameter, developed most conspicuously
near the outer surface of the coat. These patches also may have a thin fibrous core
fed by fibres visible near the periphery on the flat surfaces of the loculicidally frac-
tured carpels (PI. 12, fig. 30 ; PL 13, fig. 35). Here loops are seen from which
branches are directed inwards to the centre of the endocarp, while more numerous
and finer branches are directed outwards to its external surface.

V.i 2985 had laid long exposed before fossilization, and the coarse parenchyma
patches have decayed more readily than the more compact tissue which surrounds
them. Their decay has produced deep funnel-shaped cavities with circular orifices
which form a conspicuous feature at the surface of the endocarp. They are best
shown on the lower surface, where they occur in more or less longitudinal rows
(PI. 13, fig. 32). The whole surface of this specimen (as preserved) has been highly
polished by the abrasion it has undergone.

(3) The innermost carpellary coat is a smooth locule-lining formed of small cells
arranged so as to produce straight or slightly sinuous or criss-cross lines or striations
with a general transverse or oblique orientation. The distance between the striae
is about 0-02 mm. The striate lining is seen to the left of the axis in PI. 13, fig. 36,
and is more clearly visible on the specimen itself and on the surface of the abortive
locule-cast.

There is some reason to think that that part of the thick parenchymatous coat
which was in contact with the locule-lining was rather spongy in texture, or that
it was at least softer and less resistant to decay than the layers outside it. In places
it is partially decayed, while in other places it appears obscurely columnar in trans-
verse section (PI. 13, fig. 35, to the left of the fertile locule-cast). It clings in patches
to the locule-cast, giving the appearance at first sight of a warty testa formed of fine
angular, parenchymatous cells (PL 13, fig. 35 ; PL 14, fig. 41). But careful scrutiny
reveals its true nature. The decay is most conspicuous around the axis at the base,
exposing the locules and seeds in some specimens (PL 12, fig. 26 ; PL 13, fig. 32).
Length of fruit, 7-5 to 9-5 mm. ; transverse diameter, 17 to at least 22 (? 24) mm.
Length of ¥.12985, 8 mm. ; diameter, 19 mm. Length of specimen shown in

UPPER CRETACEOUS AND EOCENE FRUITS FROM EGYPT 171

PI. 14, figs. 42, 43, with exocarp preserved, 14 mm. ; estimated diameter, 24 by 17
mm. Length of another fruit, 10 mm. ; diameter, 22 mm.

Seeds (PI. 13, figs. 33, 35, 37 ; PI. 14, figs. 40, 41, 43-47 ; PI. 15, fig. 48) : More
or less obovate in outline, very slightly inflated, with a small beak-like projection
at the hilar end which carries the micropyle. This organ is indicated by the radial
alignment of the cells at its tip. That the radicle probably lay within the beak is
suggested by its shape. The general symmetry indicates an anatropous seed with
hilum in the concavity which delimits the " beak " from the main body of the seed,
i.e., closely adjacent to the micropyle. Here the funicle is seen to enter the seed
(cf. PI. 12, fig. 28 ; PI. 13, fig. 35). Raphe marginal, ventral, indicated by the form
of the seed as well as by a shallow marginal groove at the rounded end opposite
to the micropyle. The groove ends on the dorsal surface. Its termination probably
indicates the position of the obscure chalaza.

On the rounded end of the best developed seed exposed by abrasion within a
locule-cast of ¥.12985 the chalaza is probably marked by a small deep depression
in the same position (i.e., at the lowest point of the seed as it lay in the fruit). If
so this is the cast of the raphe fibres, where they turn sharply inwards to enter the
seed or albumen.

Testa apparently only one cell thick, since the cells of the external and internal
impressions appear to agree exactly in size and character. It was formed of equi-
axial polygonal angular cells about 0-025 to 0-03 mm. in diameter, convex externally
(as shown by the concave external impressions on a small fragment of external cast
seen in the specimen figured in PI. 14, figs. 42-47), concave internally (shown by
the convex impressions on the internal casts of the seed figured in PI. 14, figs. 39-
47). Dimensions of seeds : Length, 5-5 to about 7 mm. ; breadth, 4 to 5 mm. ;
thickness, i to 1-5 mm.

REMARKS AND AFFINITIES. In examining and interpreting mineralized fruits
it is necessary to remember that mineral substance in solution may percolate through
all incipient fissures as well as into actual cavities. On evaporating and hardening
it may then form a cement which prevents or hinders separation of parts along
natural planes of weakness. But in that the film of cement may often be incom-
plete, indications of natural dehiscence can usually be detected. Thus in Wether-
ellia (Reid & Chandler, 1933 : 251), while many specimens were so cemented that
their loculicidal dehiscence was concealed, the fruits were found in such large numbers
that plenty of evidence as to their natural manner of splitting was available. In
the fossil just described septicidal dehiscence may be obscured in a similar way.
The mineral cement may cause adhesion of surfaces and tissues which in life, or at
least after maceration, whether natural or artificial, would separate readily. For
example, in Palaeowetherellia, a locule-cast may simulate a seed (which it covers and
conceals), and patches of endocarp adhering to the cast but torn from adjacent
tissues may look like testa (cf. PI. 13, fig. 35 ; PI. 14, fig. 41). Again thin films of
cement between delicate tissues may reproduce as external or internal impression
coats which could scarcely have escaped destruction, e.g., the delicate one-cell
thick testa of W ether ellia and Palaeowetherellia.

Palaeowetherellia is now known from more than a dozen specimens. In addition

172 UPPER CRETACEOUS AND EOCENE FRUITS FROM EGYPT

to those figured by Heer (1876) as Diospyros, ten more, five of which are incomplete,
are now available. These will be catalogued below with details of any special
features they show and a note of the place of origin.

The best preserved for study was described and figured by Krausel (1939 : 106,
pi. 2, figs. 9, 10 ; text-fig. 32) as Diospyros schweinfurthi Heer. Certain of its
characters, including the number of locules, were obscure at the time, but have
since been clearly shown by the beautiful section (largely a natural fracture surface)
now exposed (cf. PL 13, figs. 33-37). On this section most of the newly recorded
details are based. Two other specimens from the Kharga Oasis have yielded addi-
tional data, thanks to their broken condition, and have made it necessary to unite
Diospyros schweinfurthi Heer and Royena desertorum of Heer as a single species
(cf. p. 176). Other fruits or fragments from Kosseir throw light on the variation
in the number of locules, the branching of fibres of the endocarp, mode of dehiscence,
variations of size, and character of exocarp.

The new evidence makes it impossible any longer to refer this species to the
family Ebenaceae in spite of some superficial resemblances to Diospyros. In this
genus the soft pulpy fruit breaks irregularly ; its radially arranged locules (with
coarse striae) have no flattened area where the two opposed surfaces are in contact.
The seeds are pendulous on short funicles which spring from the upper inner angles
of the locules arising at the top of the fruit axis (Text-fig, i A) . The anatropous seeds
have a conspicuous superficial dorsal marginal raphe which does not terminate at
the chalaza (i.e., at the opposite end of the seed to the hilum) but is continued along
the ventral margin so as to encircle the seed. It gradually tapers, dying out finally
close to the hilum. The seeds lie vertically in the locules with their ventral margins
parallel with and close to the axis. They are not beaked like those of the fossil
but the terminal micropyle usually leads into a large conspicuous canal produced
by incurving of the testa. The radicle of the embryo occupies this canal, which
on an internal cast of the seed would appear as a conspicuous deep depression or
pit (Text-fig. IE) with a projection from its base representing the cast of the radi-
cular pocket. Similar features are characteristic of Royena, which resembles the
fossil even less than Diospyros in the form and fewness of its seeds. One other
characteristic may be noted, namely the cell-structure. In Diospyros the testa cells,
while they may be similar in style to those of the Egyptian fossil, are convex intern-
ally, so that on an internal seed-cast they would produce concave instead of convex
impressions. Unfortunately it has not yet been possible to find any living closely
allied genus, although a suggestion will be made below as to family relationship.
But there can be no doubt at all that Palaeowetherellia closely resembles the fossil
genus W ether ellia Bowerbank from the London Clay in its form, placentation,
locule characters, seeds, and cell-structure, and in the tendency for dissolution of
the carpel to occur at the centre which allows the seeds to escape through the gaps
thereby produced — a point noted by Bowerbank in describing Wetherellia. Similar
decay may be seen in crab apples which have lain long on the ground in winter.
A detailed description of Wetherellia variabilis was given by Reid & Chandler (1933,
251, pi. 9, figs. 7-22). A second species — W. dixoni (Carruthers) — has since been
recognized from the Cuisian (?) — or it may be Lutetian or Auversian — of Selsey.

UPPER CRETACEOUS AND EOCENE FRUITS FROM EGYPT

173

Its full description awaits publication. The characters of these two species of
Wetherellia are summarized in a table below.

W ether ellia variabilis Bowerb.
Fruit :

A syncarpous 2 to 5-loculed septicidal
capsule or schizocarp, later splitting locu-
licidally. Sub-globular to ovoid with length
equal to, greater, or less than the diameter.
Smooth, ribbed, or angled externally.

Locules :

Radially arranged around a central axis
which extends throughout the length of the
fruit. Tangentially compressed so that the
opposed walls are contiguous except where
the seed lies.

Placentation :

Axile ; solitary seeds suspended on long
arched funicles which spring from the axis
at a point about one-third or one-quarter
of the length of the fruit from the apex.
The funicles lie within that part of the locule
where the opposed walls are contiguous.

Seed:

Slightly oblique, with the distal end nearer
to the axis than the proximal.

Pericarp :

Of thick angular parenchyma. Locule-
lining smooth and shining, obliquely or
transversely and finely striate.

mm. Diameter, 12

Dimensions of fruit :

Length, 12 to 20
to 24 mm.

Seed :

Scarcely inflated, elongate obovate in out-
line ; slightly beaked at the narrow hilar
end, anatropous ; hilum terminal, raphe
ventral, micropyle adjacent to the hilum ;
chalaza small and inconspicuous at the
opposite end to the hilum.

Testa thin, a single layer of angular equi-
axial cells convex externally, concave intern-
ally, 0-03 to 0-05 mm. in diameter.

Dimensions of typical seeds :
12 X 3-75 X 1-5 mm.

Wetherellia dixoni (Carr.)

A syncarpous 5 to 7 (or more ?) loculed
capsule splitting septicidally and loculici-
dally. Sub-globular but somewhat dorsiven-
trally compressed. Longitudinally ribbed
externally. Septal fibres apparently give
rise to hollow external spines.

As for W. variabilis.

As for W. variabilis except that the
funicles spring from the axis at a point
about one-sixth to two-sevenths of the length
of the fruit from the apex.

As for W. variabilis, but the seeds some-
what broader in proportion to their length.

As for W. variabilis.

Length, 4 to 7 mm. Diameter 13-5 to 15
mm. (somewhat dorsiventrally crushed).

As for W. variabilis, but somewhat shorter
and broader. Raphe fibres not actually seen.

Testa not seen.

(1) 5-5 x 4 x i mm.

(2) 7 x 4-75 X 1-5 mm.

UPPER CRETACEOUS AND EOCENE FRUITS FROM EGYPT

It may be noted that the parenchymatous tissues of both Wetherellia and of the
Egyptian fruits is evidently very liable to decay, for partial destruction of the centre
of the fruit has occurred in both so that the locules are exposed, allowing the seeds
to escape. Consequently locules which have not dehisced in the normal way may
be empty.

While therefore Wetherellia and the Egyptian fossils show a striking measure of
agreement such as suggests a family relationship, their characters are sufficiently
distinctive to indicate a generic difference between them. The name Palaeowether-
eUia is here suggested for the Egyptian fruits, which thus become Palaeowetherettia
schweinfurthi (Heer).

The distinctions so far as they are known at present may be summarized in tabular
form.

Wetherellia

Septicidal capsule or schizocarp, also
splitting loculicidally. Locules 2 to 7 (or
more ?).

Long .arched funicles springing from the
axis at one-sixth to one-third of the length
of the fruit from the apex.

Seed lying slightly obliquely in the locule
with the distal end nearer to the axis than
the proximal.

Seeds scarcely inflated, oval to elongate
obovate in outline, beaked at the narrow
end. Hilum terminal.

Palaeowetherellia

Loculicidal capsule also splitting septi-
cidally.

Locules 6 to about 12.

Long arched funicles springing from the
axis at one-third to one half the length of
the fruit from the apex.

Seed so obliquely placed as to be almost
transverse, with the proximal end towards
the outer edge of the fruit and the distal end
near the lower end of the axis.

Seeds slightly inflated, obovate in outline,
beaked at the narrow end. Hilum almost
terminal just below the beak on the ventral
margin.

In 1933 Reid & Chandler referred Wetherellia to the family Linaceae but it is
necessary to correct this ascription on the following grounds :

(1) More than five locules are now known to occur in undoubted Wetherellia
(W. dixoni}.

(2) As noted in 1933 the coat of Hugonia (Linaceae) with which Wetherellia was
compared is fibrous, that of Wetherellia and of Palaeowetherellia is consistently
parenchymatous with a few scattered fibres.

(3) The seed in Hugonia occupies more of the locule than the seed of Wetherellia
or Palaeowetherellia.

(4) The funicle is short and straight in Hugonia, not long and arched as in
Wetherellia and Palaeowetherellia.

(5) Polygonal cells of the testa are much finer in Hugonia than in Wetherellia or
Palaeowetherellia .

(6) The chalaza of Hugonia forms a large conspicuous scar unlike the inconspicu-
ous scarcely distinguishable chalaza of Wetherellia and Palaeowetherellia.

Taken together these differences appear on maturer reflection to be such as to
distinguish Wetherellia and Palaeowetherettia from Linaceae.

UPPER CRETACEOUS AND EOCENE FRUITS FROM EGYPT 175

The true relationship of the fossils therefore remains to be discovered. They
may belong to an extinct family whose nearest living allies have not yet been traced.
But certain features point to Euphorbiaceae as a possible alliance. These features
are the combination of loculicidal and septicidal dehiscence common in capsules of
Euphorbiaceae, the number of radially arranged locules, and their tangential com-
pression— such an arrangement is present in Hura — and the point of origin of the
funicles from the axis which may be well below the apex of the fruit in many Euphor-
biaceae. Pendulous seeds solitary in the locules with ventral raphe. At the same
time it must be frankly admitted that no really comparable genus has yet been dis-
covered, so that attribution to Euphorbiaceae must be regarded as doubtful. But
no other known family shows so many of the characters of the fossil as the Euphor-
biaceae.

MATERIAL.

Fruit (PI. 13, figs. 31-37). Also figured Krausel (1939, pi. 2, figs. 9, 10 ; text-
fig. 32). Neotype.

A nine-loculed fruit now fractured longitudinally to show one fertile and one
abortive locule, the median axis and mode of placentation.

Cell patches of the inner carpellary layers adhere to the fertile locule-cast giving
it a roughened granular appearance which at first sight simulates a rough testa
(PI. 13, figs. 33, 35). The elongate cells or striations of the locule surface are visible
on the abortive locule-cast and the locule wall between it and the axis (PI. 13, figs.
34, 36). The base of the specimen has decayed probably through long exposure
before fossilization, displaying locules, locule-casts and seeds (PI. 13, figs. 32-34).
The carpel wall has also suffered from differential decay, which has produced deep
superficial pits originally occupied by coarse-celled tissue especially on the lower
surface (PI. 13, fig. 32). Incipient loculicidal splitting is visible on one radial rib
at the apex of the fruit (PL 13, fig. 31). Diameter, 19 mm. ; height, about 8 mm.
(somewhat reduced by decay at the base).

From the Lower Danian (Upper Cretaceous) ; Farafra, Egypt. Brit. Mus. (N.H.),
No. ¥.12985.

Part of a fruit (PL 13, fig. 38 ; PL 14, figs. 39-41)-

A wedge-shaped loculicidal segment of a fruit bounded by the external surface
on one side, and by two adjacent locules on its two lateral faces. The inner end
of the wedge (fruit wall and axis) has broken or decayed, leaving exposed the locule
and seed-casts projecting inwards (PL 14, fig. 40). Tangential breadth of segment,
6-5 mm. ; height, 7-5 mm. Radius from centre (as preserved) to outer edge, 7-5
mm. Estimated breadth of fruit, about 17 mm. Length of seed or locule-cast,
5-5 mm. ; breadth, 4 mm. ; thickness, i mm.

The exocarp is not preserved, the surface of the capsule is rough, not differentially
decayed before fossilization as in ¥.12985. The substance is parenchyma with
cells about 0-02 mm. in diameter in which patches of coarser parenchyma (cells

176 UPPER CRETACEOUS AND EOCENE FRUITS FROM EGYPT

about 0-05 mm. in diameter) are embedded. The decay of such coarse patches
produced the deep pits in ¥.12985.

The locule-casts narrow to the exterior and are directly slightly upwards, at first
sight suggesting parietal placentation. One was more fully displayed by removal
of a few adherent fragments of the opposed carpel wall. Some of the parenchyma
cells of the wall adhere at this narrow end of the locule-cast producing the effect
of a rough nodular testa (PI. 14, fig. 41).

Nearer the inner end of the locule abrasion has removed first the parenchymatous
cells of the carpel and then a layer of locule-cast, thereby exposing the seed-cast
(PI. 14, fig. 41 sc.), with its angular equiaxial convex cell-impressions about 0-025
to 0-03 mm. in diameter (hence cells were concave inwards on the testa).

The testa is also represented by fragments of its external impression showing the
same cell-impressions which are concave on this surface (hence the external surface
had convex cells).

This fruit segment is clearly identical in character and size with Heer's figure of
Royena desertomm (1876, pi. i, figs. 11-16). But the details of its cell-structure
and the arrangement of its locules also unite it with ¥.12985 and with another
specimen figured in PI. 14, figs. 42-47 ; PI. 15, fig, 48. Hence this imperfect fruit
constitutes a most important link in the evidence which unites Royena desertorum
with Palaeowetherellia schweinfurthi (Heer) .

From the Dano-Montian Lower Esna Shales ; Gebel Um-el-Ghanaim, Kharga
Oasis, Egypt. Brit. Mus. (N.H.), No. ¥.31114.

A perfect six-carpelled endocarp (PI. 12, figs. 24, 25) with base intact. The locu-
licidal sutures are clear, the septicidal more obscure except where abrasion has
removed the outermost layers. The whole upper surface has been somewhat abraded
showing the radiating groups of fine parenchyma. Diameter, 17 mm. ; length,
9-5 mm. Brit. Mus. (N.H.), No. ¥.31110.

A seven-partite endocarp, slightly asymmetrically developed. It is somewhat
corroded on one side so that a seed (or locule-cast) is partially exposed. The exo-
carp is almost entirely abraded, one small patch only persisting at the base. The
surface of the endocarp is also much abraded. The whole specimen is encrusted
with mineral deposit which forms small pimples over the exposed fibre ends and
coarse cell patches. Diameter, 17 mm. ; length, 10-5 mm. M. I. Youssef Collec-
tion, 1952.

Two segments of an endocarp showing the cavities of two locules. The lines of
loculicidal dehiscence can be detected. The surface is somewhat abraded, exposing
the irregular rugosities due to the radiating clusters of cells just below. A seed-
cast in the locule between the two segments shows clear evidence of the ventral
raphe. M. I. Youssef Collection, 1952.

The above three specimens are from the Dano-Montian Lower Esna Shales ;
Gebel Atshan, Kosseir Area, Red Sea.

An eight-loculed endocarp (PL 12, fig. 26), perfect except for decay at the centre
of the base so that the inner angles of the locules are exposed. The septa show

UPPER CRETACEOUS AND EOCENE FRUITS FROM EGYPT 177

median planes of weakness as for septicidal dehiscence. Five locules have retained
their seeds, some of which are abortive ; from the others the seeds have fallen
through the gap caused by the decay above described. No exocarp is preserved,
and there is considerable mineral incrustation over the surface of the abraded endo-
carp. Hollows for the fibre ends are visible on the lower surface ; a few encrusted
rounded knobs probably indicate their position on the upper surface.

Half of a fruit (PI. 12, figs. 27, 28) which has been fractured longitudinally through
two of the locules. One has a locule-cast preserved, in the other the locule-cast is
missing, but the funicle (now much encrusted with a mineral deposit) is seen. There
appears to be evidence of seven locules on this fragment, so that the complete fruit
must have had at least eleven or twelve locules. Adherent remains of exocarp are
seen at the base and apex only and are shown in section where the coat has broken
away from the endocarp. Exocarp and endocarp are much encrusted by a mineral
deposit. On the endocarp the encrusted remains of the fibres described on p. 170
form small rounded prominences.

Half of an endocarp which has split longitudinally (PL 12, figs. 29, 30). Three
and a half carpels are preserved and the septicidal and loculicidal fracture lines are
clearly visible.

The half carpel has split both loculicidally and septicidally and has been pushed
out of position. The surface is sufficiently abraded to expose the radial grouping
of the small cell-clusters. The network of fibres is obvious on one loculicidal suture
plane, although owing to the adherent parenchyma and secondary incrustation the
fibres are not themselves exposed but their position is very apparent. No exocarp
is preserved. Diameter of endocarp, 18 mm. ; length, 9 mm. A few small shell
casts and impressions adhere to one fracture plane.

The above three specimens are from the Dano-Montian Lower Esna Shales ;
Abu Tundub, Kosseir Area, Red Sea. Brit. Mus. (N.H.), Nos. ¥.31111-13.

A fruit (PI. 14, figs. 42-47 ; PL 15, fig. 48) with exocarp preserved but cracked
in such a manner that at first sight it simulates three perianth segments (PL 14,
fig. 42). Actual diameter (incomplete because the specimen had been polished on
one side at p in figs. 42, 43 thus showing a tangential section), 18 x 17 mm.
Estimated complete maximum diameter (distorted), about 24 X 17 mm. Height,
14 mm.

The whole fruit is cracked through desiccation and crushing, and is held together
by calcite cement (white in the figures). On the polished surface two seeds can be
seen, one in transverse and one in oblique section. The first shows the seed-cast
(sc) surrounded fairly closely by the locule-cast (Ic) (PL 15, fig. 48), which is em-
bedded in the pulp or parenchyma of the carpel. The base of the fruit is sunk,
and broken along one radius so that a beautiful seed-cast was exposed, at first
held in place by calcite cement, but later becoming detached (PL 14, figs. 43-47).
The beaked end of the compressed ovate cast was directed outwards and upwards
in the locule as in the section of ¥.12985 shown in PL 13, figs. 33, 35, 37.

178 UPPER CRETACEOUS AND EOCENE FRUITS FROM EGYPT

Length of seed-cast, 7 mm. ; breadth, 4-75 mm. ; thickness, 1-5 mm.

Surface of cast with equiaxial slightly convex cell-impressions. Their divergence
at the beaked end indicates the micropyle. A shallow marginal furrow at the
opposite rounded end of the cast dies out on the dorsal surface (close to an acci-
dental fracture line) . It indicates the inner end of the raphe, its termination marking
the site of a small inconspicuous chalaza.

From the Dano-Montian Lower Esna Shales ; Gebel Tarawan, Kharga Oasis,
Egypt. Brit. Mus. (N.H.), No. V.3HI5.

Two segments of an endocarp showing the cavities of three locules. The carpel
wall between the two segments is abraded so that the seed-cast is exposed along its
dorsal margin. When the segments are separated the chalaza can be detected at
the lowest point of the seed-cast as it lies in the endocarp.

The exterior of the endocarp is also abraded, so that in places the inner thick layer
of fine parenchyma is exposed. Remains of the outer layer with radiating cell
clusters occur hi patches, but are much obscured by mineral incrustation. Length
of endocarp, 8 mm. ; maximum breadth across the two loculicidal segments, 14 mm.

From the Dano-Montian Lower Esna Shales ; Gebel Durvi, Kosseir Area, Red Sea.
M. I. Youssef Collection, 1952.

Family ICACINACEAE
Genus ICACINICARYA Reid & Chandler, 1933

Icacinicarya youssefi n. sp.

(PI. 15, figs. 49-51)

DIAGNOSIS. Fruit and endocarp almond-shaped or sub-obovoid in outline, lenti-
cular in transverse section. External surface of endocarp with a series of more or
less discontinuous irregular rugosities, some longitudinal, others transverse or oblique.
Cells of endocarp markedly sinuous or coarsely digitate. Length of endocarp, 31
mm. ; breadth, 22 mm. ; thickness, 10 mm. Length of a fruit, 33 mm. ; breadth,
26-5 mm. ; thickness, 13 mm.

HOLOTYPE. Endocarp figured PI. 15, fig. 51. Brit. Mus. (N.H.), No. ¥.31117.

LOCALITIES AND HORIZON. Gebel Atshan and Gebel Durvi, Kosseir Area, Red
Sea ; Dano-Montian Lower Esna Shales.

DESCRIPTION. Fruit (PI. 15, figs. 49, 50) : Bisymmetric, somewhat compressed,
giving a lenticular transverse section, obovoid in outline, slightly mucronate at the
apical style, attachment at the opposite extremity to the style. Surface much
puckered as if by shrinkage of the mesocarp, but having a few rather ill-defined
longitudinal ridges which may branch or anastomose halfway up. They may be
due to fibres in the pulp or just beneath the epicarp. The actual epicarp itself is
probably missing. One margin (overlying the funicle ?) thicker than the other
with two or three marked longitudinal ridges. Whole surface with a pattern of

UPPER CRETACEOUS AND EOCENE FRUITS FROM EGYPT 179

small, rounded or quadrangular depressions, about 0-5 to i mm. in diameter, ar-
ranged in longitudinal rows. Finer structure of digitate or markedly sinuous cells.
The convergence of the ornamentation at the two ends indicates the position of style
and attachment described. Length of fruit, 33 mm. ; breadth, 26-5 mm. ; thickness,
13 mm.

Endocarp (PI. 15, fig. 51) : Similar in form to the fruit, base narrowed to a point
where the opening for the funicle is clearly seen. A ridge due to splitting followed
by mineral infiltration is seen at the apical style. This tendency to split at
the stylar end has been observed in other Icacinaceae, e.g., Natsiatum eocenicum
Chandler from the Lower Headon of Hordle.

As in genera of Icacinaceae one margin is much thicker and more rounded than
the other. Experience has shown that the thick margin carries the funicle but no
section is available, so this cannot be verified in the present instance. External
surface with an obscure pattern of small mostly discontinuous rugosities, some
elongate and longitudinally aligned. They do not produce a definite network of
ridges and hollows as in Icacinicarya platycarpa Reid & Chandler (1933 : 345, pi.
16, figs. 11-18).

Surface of endocarp formed of small digitate or conspicuously sinuous cells with
a tendency to transverse alignment. They are about 0-114 nim. broad and 0-057
mm. in length. Length of endocarp, 31 mm. ; breadth, 22 mm. ; thickness, 10 mm.

REMARKS AND AFFINITIES. Two specimens which by their general similarity
may be presumed to belong to the same species although one shows the whole fruit,
the other the endocarp only. One surface of the fruit shows the outline of a closely
comparable endocarp owing to the contraction on drying which the pulpy exocarp
has undergone (PI. 15, fig. 50).

Although the most conclusive diagnostic characters of Icacinaceae can only be
inferred because the unbroken character of the specimens conceals them, there can
be little doubt of the relationship. The specimens are considerably larger than
Icacinicarya platycarpa, which they resemble in their form, but as pointed out
above, they can also be distinguished from that species by surface ornamentation.
The cells of the endocarp are considerably larger than in that species and differ in
their digitate form.

Although further information about this species is much to be desired, it is suffi-
ciently well defined to be recognizable again, so that the specific name, Icacinicarya
youssefi, after the finder, has been given.

Family ICACINACEAE ?

ICACINICARYA sp. ?

(PI. 15, figs. 52-54)

LOCALITY AND HORIZON. Gebel Atshan, Kosseir Area, Red Sea ; Dano-Montian
Lower Esna Shales.

DESCRIPTION. Fruit : Sub-ovoid, bisymmetric about a plane which includes the
two major axes and probably the funicle, although this was not actually seen,

GEOL. II, 4. 13

i8o UPPER CRETACEOUS AND EOCENE FRUITS FROM EGYPT

somewhat sharply angled in the plane of symmetry on one side, rounded and much
inflated on the other (funicular margin). Style terminal at the apex, marked by a
slight prominence. Attachment at the opposite end of the major axis to the style
where the fruit shows a slight flattening, marked clearly by a small circular scar
from which the surface cells radiate. The broad margin of the fruit is semicircular
in outline ; the narrow margin is slightly concave immediately below the style and
for about one-third of the length, but is convex for the lower two-thirds. Surface
puckered and wrinkled giving a leathery appearance. Surface cells obscure except
around the attachment where they are equiaxial, rounded, slightly convex, and
about 0-038 mm. in diameter. Length of fruit, 13 mm. ; breadth in plane of sym-
metry, 11-5 mm. ; thickness at right angles to plane of symmetry, n mm.

REMARKS. The solitary specimen looks like a drupe with epicarp and mesocarp
preserved so that all the characters of the endocarp are hidden except so far as they
can be inferred from the form and symmetry of the specimen. The peculiar asym-
metric outline as viewed at right angles to the plane of symmetry and the occurrence
of one broad rounded and one angled margin may indicate a fruit of Icacinaceae.
But pending the discovery of further evidence the specimen can only be referred
tentatively to the form-genus Icacinicarya.

Family FLACOURTIACEAE ?
Genus THIEBAUDIA nov.

DIAGNOSIS. A large berry with about 36 parietal placentas, numerous seeds in
two close set rows on the placentas, and a pulpy mass of tissue which occupies the
whole of the fruit cavity. Seeds probably sub-ovoid. Length of fruit (compressed),
13 mm. ; breadth (much increased by compression but bereft of pericarp), 37 mm.
Diameter of seeds, I to 1-5 mm.

TYPE SPECIES. Thiebaudia rayaniensis n. sp.

Thiebaudia rayaniensis n. sp.

(PI. 16, figs. 58-63)

DIAGNOSIS. As for genus.

HOLOTYPE. A single fruit with most of the pericarp removed. Thiebaud &
Robson Collection (Locality no. 604), 1951. Brit. Mus. (N.H.), No. ¥.31120.

LOCALITY AND HORIZON. Wadi Rayan, Western desert of Egypt ; Eocene
(? Lutetian or slightly younger).

DESCRIPTION. Fruit : A globular or sub-globular berry (now much compressed
dorsiventrally, so that the wall has been buckled all around the equator of the
specimen). Dehiscence probably by irregular breaking of the pericarp as no indi-
cation of sutures for regular dehiscence can be seen. The pericarp is preserved only
at the extreme base and apex (PI. 16, figs. 58, 59), elsewhere it has been broken away
or abraded.

The external surface shows a few obscure radial flutings at the upper end, but is

UPPER CRETACEOUS AND EOCENE FRUITS FROM EGYPT 181

preserved in a matrix so coarse that it does not show the cell structure of the epicarp
if still present.

A slightly denned and very slightly sunk area at the apex, about 5 mm. in diameter,
may mark the base of the style. Thickness of pericarp about 3 mm. Numerous
broad, rather flat, stout, longitudinal bands of fibres can be seen (although all are
now incomplete) in the lower half of the fruit lying in fragmentary remains of the
pericarp, but above the equator they appear to have been abraded. Placentation
parietal, the seeds arranged in two close-set rows on each of thirty-six longitudinal
placentas. Sometimes the seeds in the two rows are opposite one another, sometimes
they appear to be alternate. Between each pair of placentas there is a very slight
longitudinal ridge which thickens at its extreme apical end. These ridges produce
on the inner surface of the pericarp a series of broad shallow longitudinal channels
or pockets into which the placentas and seeds fitted.

Each placenta begins to produce its seeds at about 10 mm. from the apex of
the fruit, where it springs from a pointed tongue of tissue with a narrow median
furrow (PI. 16, figs. 58, 60). The whole of the interior of the fruit is filled by a pulpy
mass which adheres closely to the pericarp. The shallow ridges of the pericarp
give rise to grooves upon the surface of the pulpy-mass. These separate the broad
flat-topped ridges opposite the placentas bearing the concavities caused by the seeds.

It is the surface of this mass which is exposed intact on the upper surface of the
specimen (PI. 16, figs. 58, 60). The pulp is formed of coarsely and deeply sinuous
cells, and throughout its thickness there are numerous small cavities about the same
size as these cells (i.e., about 0-057 to 0-114 mm. in diameter.) No tendency to split
either radially or otherwise has been detected in the pulpy mass.

Length of fruit (much reduced by dorsiventral compression), 13 mm. ; breadth
(correspondingly increased by compression but reduced by the loss of the pericarp),
37mm.

Seeds : Very obscure. Producing sub-circular or sub-ovoid hollows on the pulpy
mass (PI. 16, figs. 58-61). Perhaps somewhat laterally compressed. The shallow
convex external surface of the seed was ornamented with large inflated radially
arranged cells or areoles diverging from a knob-like projection (PI. 16, fig. 62).
This structure is visible on the impression of a detached seed (probably belonging
to the fruit) which is preserved on the pericarp near the base. Obscure traces of
similar cells were also seen on a few seed-impressions on the pulpy mass. Other
seed-impressions merely show a rounded prominence which must represent a con-
siderable depression (hilar, micropylar, or chalazal ?) on the actual seed surface
(PI. 16, fig. 61, best shown in seed s). Internal casts of seeds (preserved in a few
instances on the underside of the fruit but always incomplete, PI. 16, fig. 63)
show a smooth shining surface with longitudinal striations due to very long narrow
cells with stout longitudinal walls, 0-014 to 0-018 mm. broad, lying parallel with
the long axis of the seed. There are also slight traces of a coat of transversely
aligned cells or fibres. Diameter of seeds, i to 1-5 mm. An internal cast measured
1-5 mm. in length, i mm. in breadth.

REMARKS. One fruit preserved in a ferruginous cement filled with quartz grains
forming a hard mass. It appears to have been embedded in a cream-coloured fine

182 UPPER CRETACEOUS AND EOCENE FRUITS FROM EGYPT

sand with Nummulites. The sand filled every crack and cranny of the cast, and a
Nummulite (reported by Mr. C. D. Ovey as having a range from Lutetian to Middle
Oligocene) was found in sand lying in the deep hollow between the lower surface
of the pulpy mass and the remains of the pericarp at its base. There can be little
doubt that the Nummulite belonged to the strata in which the fruit occurred, for
little attempt had been made to clean the fossil of the adherent sand which clung
with persistence. It has now been boiled and the sand brushed away. Other
fossils from these beds are also said to be ferruginous, but this is the only plant as
yet discovered. The age of the deposit is regarded as Lutetian, or possibly some-
what younger, but definitely Eocene.

As stated, most of the pericarp of this fruit was missing, revealing the central
pulpy mass. It is a difficult specimen to understand and interpret. But probably,
if its living counterpart were discovered, the description given above could be con-
siderably simplified and shortened.

The coarseness of the matrix is not well calculated to preserve fine details. Little
can be seen of the seed structure, especially as the hollows on the central mass are
largely merely cavities in the pulp due to the pressure exercised by the growing seeds.
They do not therefore give much information apart from the size and, rather ob-
scurely, the form of the close-set seeds which are themselves almost entirely absent.
Even these impressions are partially confused by some measure of overlap. The
bases of the stout fibres in the fruit wall persist in broken ragged fragments at the
base of the specimen, where they project from the fragment of pericarp. Sometimes
they are missing, and the openings from which they came can be seen in the edge of
the broken wall. The upper side of the specimen shows in good condition the
surface of the pulpy mass with impressions made by the seeds on the ridges opposite
the placentas, and the shallow furrows between these ridges which are the impressions
of the narrow ridges on the pericarp wall described above.

The lower surface is less well preserved. It appears to have become torn and
battered before or during fossilization, so that the actual lower surface of the pulpy
mass together with the alignment and ridges on this surface is largely destroyed.
Traces only of these features and of the collapsed and fragmentary pericarp wall
with its fibres can be detected. Detached seeds represented by occasional external
impressions or true internal casts have been pushed into the soft pulp and are visible
here and there. Usually the casts are obscure. The true internal casts are rarer
than the external impressions.

A curious and at first misleading feature of this specimen is a coarse transversely
elongate meshwork of angular ridges around the equator. A radial fracture of the
pulpy mass demonstrates the purely secondary and inorganic character of these
ridges, which are due to the infiltration and setting of a limonite cement in cracks
caused by buckling of the walls and pulp. No organic structure is to be seen in the
seams of limonite which fill these cracks.

AFFINITIES. The parietal placentation of this multi-carpelled fruit limits relation-
ship to very few families. At first sight the specimen recalls a Poppy capsule on
account of its radial symmetry and rounded small seeds, while the remains of the
fruit wall at the apex simulate the stigma-bearing disc of Papaver. Here, however,

UPPER CRETACEOUS AND EOCENE FRUITS FROM EGYPT 183

the resemblance ends. The pulpy mass of the interior is wholly unlike anything in
Papaveraceae. Parietal placentas associated with such a mass of pulp do, however,
occur in Flacourtiaceae. But the difficulty here is that in no known genus of that
family do as many as 36 placentas and carpels occur, while the details of seed structure
in the fossil are too imperfectly known to be conclusive. No other living family
appears to bear so close a relationship to this lovely fossil, and past experience has
demonstrated the tendency for a larger number of locules to occur in fossil than in
living forms, although not perhaps to the degree here recorded.

In the absence of more satisfactory information the specimen has been referred
doubtfully to the family Flacourtiaceae. It has been given a new generic name,
Thiebaudia, after one of the finders, while the specific name rayaniensis indicates the
place where it was found.

The writer is of opinion that this specimen ought to be compared very carefully
with Krausel's species Nymphaeopsis lachmanni from the Lower Oligocene of Cairo.
The supposed placenta and unequal hammer-shaped involucral segments of that
species may quite possibly be parts of a pericarp which has cracked and contracted
into irregular segments on drying. Such a feature is displayed by one fruit of
Palaeowetherellia from Kharga (PI. 14, fig. 42), in which the cracked berry with its
mineral infiltrations simulates a trifid perianth. The fibres shown by Krausel
(1939, pi. 2, figs. 2, 3, 6) on the surface exposed beneath the supposed involucral
segments are not altogether unlike those of Thiebaudia. The direction in which
they branch suggests that the supposed apex is in fact the base of the specimen.
The buckling of the fruit at the equator due to dorsiventral compression has pro-
duced a similar network of infiltration ridges to those described in Thiebaudia.
It is not impossible that the apparent operculum of the seeds might be interpreted
as a large chalazal scar. But these are merely suggestions for future consideration.
They cannot be substantiated without the most careful study of the specimens
themselves.

Carpolithus hassani n. sp.
(PI. 16, figs. 64, 65)

LOCALITY AND HORIZON. Gebel-el-Ter, Kharga Oasis, Egypt ; Dano-Montian
Lower Esna Shales.

A beautiful but puzzling specimen suggests one valve of a bisymmetric endocarp.
It appears to have been rubbed down artificially at the margins but to no great
depth. The outline, as it exists at present, is broadly elliptical, 20 mm. long,
17-5 mm. broad, 5-5 mm. deep. The external surface is gently convex and the
internal correspondingly concave. The thickness of the wall at the polished margin
is 2 to 2-5 mm.

Three deep external grooves must have some structural significance. They
appear to be the outer edges of slits which pass through the whole thickness of the
wall. The two nearest the margins of the valve slope inwards towards the centre
of the fruit so that their inner edges, marked by deep grooves on the surface of the
locule, lie slightly closer together than do their outer edges on the external surface

184 UPPER CRETACEOUS AND EOCENE FRUITS FROM EGYPT

of the valve. These two grooves are parallel with the outline of the valve, but
they do not meet, although their ends lie much closer together at one extremity
(apex ?) of the valve than at the other. One groove is longer than the other. The
third groove which pierces the endocarp also, lies midway between the other two
but is shorter than either. It begins at about the middle of the valve and passes
towards that end of it (base ?), where the other two grooves are most widely separated.
It dies out before reaching the edge of the valve. Within the slits the surface of
the endocarp is longitudinally striate.

On the exterior, between the longer curved marginal groove and the short median
one, a longitudinal slightly sinuous furrow can be seen which was evidently a channel
for a fibre, impressions of which are seen. It gives off two or three short slender
branches at its upper end. These diverge and taper upwards.

Surface of valve rough, due to the convex angular walls of the parenchymatous
cells of which it is composed. Cells 0-05 to o-i mm. in diameter. Locule surface
smoother, formed of equally large equiaxial cells.

A first glance at this specimen suggests that it may belong to Menispermaceae.
But closer scrutiny shows that the relationship is impossible for the following reasons :

(1) The curved marginal grooves do not unite at the apex to form a horseshoe,
nor does one of their opposite ends curve appreciably outwards.

(2) The external grooves correspond with internal grooves and not with internal
ridges as in Menispermaceae, where the ridges form a cavity for the curved seed.

(3) The endocarp of Menispermaceae is fibrous in structure, not parenchymatous.

I have found no fruit with such peculiar slits which may be connected with ger-
mination. Possibly the curved area enclosed by the slits may be associated with a
curved embryo.

Carpolithus sp. (Icacinicarya sp. ?)
(PI. 16, fig. 66)

LOCALITY AND HORIZON. Gebel-el-Ter, Kharga Oasis, Egypt ; Dano-Montian
Lower Esna Shales.

The internal cast of a valve of an endocarp 10-5 mm. long, 9-25 mm. broad, 1-8
mm. deep. The cast is formed of coarse crystalline ferruginous matter. Its surface
shows an obscure network of ridges. The internal surface of the actual specimen
would have shown corresponding grooves separated by shallow convexities. No
cell-structure is visible. The form of the valve suggests Icacinaceae, but the evidence
preserved is insufficient for certainty.

Carpolithus sp.

(PL 16, figs. 55-57)

LOCALITY AND HORIZON. Gebel Atshan, Kosseir Area, Red Sea ; Dano-Montian
Lower Esna Shales.

A sub-globular fruit slightly laterally compressed, with a three-angled apex, the
angles meeting in a slight prominence. They die out at about the middle of the
fruit. Between these well-marked angles a few subsidiary inconspicuous longitu-

UPPER CRETACEOUS AND EOCENE FRUITS FROM EGYPT

185

dinal ridges can be seen. Basal attachment small, slightly sunk, marked by a
little projection at the middle of the hollow. At the extreme base of the fruit there
are three short, very shallow, rounded furrows opposite the three apical ridges or
angles. Surface, as now preserved, ornamented with numerous angular, more or
less equiaxial contiguous concave areas, somewhat variable in size, up to I or 1-5
mm. in diameter but sometimes smaller. This cast is formed of radiating groups
of fine cells, the groups often but not invariably coinciding with the concavities.
Length of fruit, 13-5 mm. ; diameter, 10 by 13-5 mm.
There is nothing to indicate the identity of this specimen.

APPENDIX I

NOTE ON THE OCCURRENCE OF THE FOSSIL FRUITS AND SEEDS
COLLECTED FROM THE KHARGA OASIS, WESTERN DESERT

OF EGYPT

By M. YOUSSEF HASSAN

The specimens were collected from a bed with an average thickness of 35 metres.
It is composed of dark grey and greenish shales, well foliated and often densely
seamed with gypsiferous and salt intercalations. Red nodules of botryoidal limonite,
sometimes attaining large sizes, are abundant. Frequently the limonite appears
in pseudocrystalline form, being cubes sometimes with interpenetration twinning.
The bed is rich in fossils which are dwarfed and excellently preserved in limonite.
The age of this bed is " Danian " or more probably Dano-Montian. The following
is a complete list of the fossil fauna identified :

Schizorhabdus libycus Zittel.

Palaeopsammia multiformis Wanner.

Bathypsammia cleopatrae Hassan MS.

Trochocyathus epicharis Wanner.

Trochocyathus deniseptatus Hassan MS.

Dungulia libyca (Wanner).

Pattalophyllia aegyptiaca (Wanner).

Terebratulina sp.

Pentacrinus sp.

Nucula tremolate-striala Wanner.

Nucula chargensis Quaas.

Leda leia Wanner.

Trapezium sp.

Trapezium aff. acutangulum (Deshayes).

Cardium cf . becksi Muller.

Cardium cf. inaequiconvexum Cossman &

Pissarro.

Crassatella zitteli Wanner.
Cucullaea schweinfurthi Quaas.
Cucullaea sp.
Limea sp.

Chlamys mayer-eymari (Newton).
Verticordia nova Hassan MS.
Corbula striatuloides Forbes.
Trochus cf. laryi D'Archiac & Haime.

Scala calamistrata (Wanner) .

Architectonica dachelensis (Wanner).

Natica (Gyrodes) farafrensis Wanner.

Natica (Euspira) terensis Hassan MS.

Rissoa cf. crassistriata Wood.

Campanile cf. brookmani Cox.

Cerithium abictiforme Wanner.

Cerithium bigeniculatum Wanner.

Cerithium cf. periphractum Wanner.

" Alaria " schweinfurthi Quaas.

Cypraea cf. kayei Forbes.

Tonna sp.

Tudicla peroni Quaas.

Sassia tuberculosa (Kaunnowen).

Sassia farafrensis (Quass).

Sassia chalmasi (Quaas).

Sassia sp.

Athleta (Volutilithes] daniensis (Quaas).

Cyclichna cf. regulbiensis (Adams).

Solidula chargensis (Quaas).

Solidula pharaonum (Wanner) .

Avellana cretacea Quaas.

Pyrgopolon sp.

Nautilus applanatus Wanner, ex Zittel MS.

Nautilus desertorum Quaas, ex Zittel MS.

1 86

This horizon is resting on a series of alternating marls and impure limestones
with occasional phosphatic bands of undoubtedly late Maestrichtian age.

APPENDIX II

NOTE ON THE OCCURRENCE OF THE FOSSIL FRUITS AND

SEEDS COLLECTED FROM THE KOSSEIR AREA

(RED SEA), EGYPT

By MURAD I. YOUSSEF

The fruits were collected from a single bed in different localities in the Kosseir
Area (Red Sea). The bed in which these fossil fruits, as well as the fossils mentioned
below, were found is a greyish-green shale varying in thickness between 50 and
125 metres. The concretions found throughout this bed, and the fossils occurring
generally in a band near its middle part, are all limonitic. Cubic pseudomorphs of
limonite, probably after pyrite, are also found. Irregular gypsum veins running
in every direction, perhaps corresponding to the more or less pyramidal planes of
jointing of the shales, are epigenetic, being secondary in origin.

The fossils collected from this bed are considered to be Danian or Dano-Montian.
Some 30 metres of shales and marls lying immediately below this bed are of the
same age. These are underlain by Maestrichtian rocks containing many phosphate
and phosphatic beds.

The following is a list of the fossil fauna identified from the thick shale bed :

Brachycyathus daniensis Wanner.
Trochocyathus epicharis Wanner.
Pattalophyllia aegyptiaca (Wanner).
Caryosmilia granosa Wanner.
Dungulia milneri (Gregory).
Dungulia libyca (Wanner).
Caryophyllia jasmundi Wanner.
Palaeopsammia multiformis Wanner.
Stephanophyllia (Microbacia) sp.
Serpula cf . discoidea Wanner.
Pentacrinus sp.
Salenia sp.
Cyphosoma sp.

Hemiaster chargensis Wanner.
Hemiaster (Leucaster) lamberti Cottreau.
Terebratulina chrysalis (Schlotheim).
Terebratulina sp.
Nucula chargensis Quaas.
Nucula lucida J. Boehm.
Nucula tenera J. Miiller (non S. V. Wood).
Nucula tremolate-striata Wanner.
Leda leia Wanner.
Thyasira cretacea (Wanner).
Cardium cf. inaequiconvexum Cossmann &
Pissarro.

Cardium cf. becksi Miiller.

Trapezium sp.

Lucina dachelensis Wanner.

Crassatella matercula Mayer-Eymar.

Area modioloides Wanner.

Cucullaea sp.

Chlamys mayer-eymari (Newton).

Verticordia nova Hassan MS.

Scala cf . desertorum Wanner.

Natica farafrensis Wanner.

Campanile cf. brookmani Cox.

Alaria schweinfurthi Quaas.

Alaria sp.

Sassia chalmasi (Quas).

Neptunea zitteli (Quaas).

Neptunea sp.

Fusus sp.

Athleta (Volutilithes) desertorum (Quaas).

Solidula chargensis (Quaas).

Avellana cretacea Quaas.

Avellana sp.

Cylichna cf. regulbiensis (Adams).

Dentalium bicarinatum Wanner.

Nautilus desertorum Quaas, ex Zittel MS.

UPPER CRETACEOUS AND EOCENE FRUITS FROM EGYPT 187

REFERENCES

AXELROD, D. I. 1952. Variables affecting the Probabilities of Dispersal in Geologic Time.

Bull. Amer. Mus. Nat. Hist., New York, 99 : 177-188, 2 figs.
BONNET, E. 1904. Sur un Nipadites de 1'eocene d'Egypte. Bull. Mus. Hist. nat. Paris, 10 :

499-502, 2 figs.
BOWERBANK, J. S. 1840. A History of the Fossil Fruits and Seeds of the London Clay.

144 pp., 17 pis. London.
CHANEY, R. W. 1940. Tertiary Forests and Continental History. Bull. Geol. Soc. Amer.,

New York, 51 : 469-488, pis. i, 2.

1947- Tertiary Centers and Migration Routes. Ecol. Monogr., Durham, N.C., 17 : 140-148.

EDWARDS, W. N. 1936. The Flora of the London Clay. Proc. Geol. Ass. Lond., 47:22-31,

3 figs.
ENGELHARDT, H. 1907. Tertiare Pflanzenreste aus dem Fajum. Beitr. Paldont. Geol. Ost.-

Ung., Wien, 20 : 206-216, pis. 18, 19.
HEER, O. 1876. Ueber fossile Friichte der Oase Chargeh. N. Denkschr. schweiz. Ges.

Naturw., Zurich, 27 : i-n, pi. i.
KRAUSEL, R. 1939. Ergebnisse der Forschungsreisen Prof. E. Stromers in den Wusten

Agyptens, IV. Die fossilen Floren Agyptens, 3. Die fossilen Pflanzen Agyptens, E.-L.

Abh. bayer. Akad. Wiss., Munchen (N.F.) 47 : 1-140, pis. 1-23.

& STROMER, E. 1924. Ergebnisse der Forschungsreisen Prof. E. Stromers in den

Wusten Agyptens, IV. Die fossilen Floren Agyptens, 3. Die fossilen Pflanzen Agyptens,
A-C. Abh. bayer. Akad. Wiss., Munchen (N.F.) 30, 2 : 1-48, pis. 1-3.

KRYSHTOFOVICH, A. N. 1929. Evolution of the Tertiary Flora in Asia. New Phytol., Cam-
bridge, 28 : 303-312.
LE MAOUT, E., & DECAISNE, J. 1876. A General System of Botany. 2nd edit. 1066 pp.,

5500 figs. London.
REID, C., & E. M. 1915. The Pliocene Floras of the Dutch- Prussian Border. Meded.

Rijksopsp. Delfst., Amsterdam, 6 : 1-178, pis. 1-20.
REID, E. M., & CHANDLER, M. E. J. 1926. The Bembridge Flora. Catalogue of Cainozoic

Plants in the Department of Geology, 1. viii -+- 206 pp., 12 pis. Brit. Mus. (Nat. Hist.),

London.
1933- The Flora of the London Clay, viii + 561 pp., 33 pis. Brit. Mus. (Nat.

Hist.), London.
RENNER, O. 1907. Teichosperma, eine monokotylenfrucht aus dem Tertiar Agyptens. Beitr.

Palaont. Geol. Ost.-Ung., Wien, 20 : 217-220.
SCHENK, A. 1889. Palaeophytologie. In Zittel, K. A., Handbuch der Palaeontologie, 2 : 958

pp., 433 figs. Munich & Leipzig.
SEWARD, A. C. 1934- An Extinct Malayan Flora in England. Sci. Progr. Twent. Cent.,

London, 29, i : 1-24.

& ARBER, E. A. N. 1903. Les Nipadites des couches Eocenes de la Belgique. Mem.

Mus. nat. Belg., Bruxelles, 2 : 1-16, pis. 1-3.

NOTE. — While this paper was going through the press my attention was drawn to a
memoir by L. W. LeRoy (Biostratigraphy of the Maqfi Section, Egypt. Mem. Geol. Soc.
Amer., Washington, 54, 1953) m which, from a study of the Foraminifera, the author concludes
not only that the " Esna Shales " are definitely Lower Eocene, but also that the Danian is
probably unrepresented in Egypt. There may, however, be some unresolved confusion over
the use of the term " Lower Esna Shales."

GKOL. II, 4 I4

PLATE 10

Nipa burtim (Brongniart)

FIG. i. The domed apical end of a drupe which is incomplete below the middle. Length
preserved, 25 mm. ; breadth, 27 mm. Estimated complete length, about 45-50 mm. It shows
the typical longitudinal fibro-vascular bundles embedded in parenchyma, (u) the apical
umbo. x 2-8. (¥.31105.)

FIG. 2. The same, viewed from below, looking on to the fractured surface. The side shown
in fig. i is towards the top of the figure. Part of the unridged smooth endocarp (e) is exposed.
Transverse fibres of the endocarp are obscurely seen. The pericarp is in section at (p, p) and
in surface view with its longitudinal fibres at (/). X 2-8.

FIG. 3. An obliquely distorted seed-cast in which the basal aperture (a) occupies a basi-
lateral position. Slight longitudinal flutings of the surface are visible. There is a mosaic
pattern all over the surface, x £ approx. (V. 13239.)

FIG. 4. The opposite surface of the same seed-cast, apical end, showing the sub-apical
funnel-shaped opening (ch) and the raphe-fibre impressions converging towards it and passing
into it. (m) marks the apical mucro of the nut. x 1-2.

FIG. 5. A seed-cast so compressed as to be almost lenticular, (a) is the position of the basal
aperture. The coarse network of fibres which lay between the two integuments of the testa
is faintly impressed upon the cast to the left of the median line. X J approx. (¥.13240.)

Nipa fruticans Thunberg

FIG. 6. A seed for comparison with fig. 5 from which the outer coat of the testa has been
entirely removed. Most of the inner coat has also been removed except beneath the slightly
sunk coarse network of fibres ( /, / ) which lay between the two coats and which still adheres
to the seed. In the meshes between the fibres (where the inner coat has gone) the transverse
alignment of the small ruminations of the endosperm can be seen rather obscurely. The seed
has been cut longitudinally. X 2 approx. Recent ; Singapore.

Bull. B.M. (N.H.) Geol. 2, 4

PLATE 10

2 f

NIPA BUKTINI, NIPA FRUTICANS

PLATE 11

a burtini (Brongniart)

FIG. 7. Opposite side of the seed-cast in PI. 10, fig. 5, showing detail of endosperm
structure. The elongate alignment at (r) indicates the position of the raphe, the radial arrange-
ment indicates the point of entry of the raphe fibres (ch) below the apex of the cast, x 1-8.

FIG. 8 The same seed-cast, detail of the surface depicted in PI. 10, fig. 5, showing the
more normal transverse alignment of the endosperm, x 1-8.

FIG. 9. The base of a seed with its ribbed outer integument partially preserved but suffi-
ciently abraded to expose the flat broad fibre band (/). The basal aperture is clearly seen.
The specimen has undergone much lateral compression, x i. Figured by Krausel, 1939,
pi. i, fig. 24 as Rubiaceocarpum markgrafi.

Nipa fmticans Thunberg

FIG. 10. A seed, side, showing the ribbed outer integument with clear impressions of the
longitudinal, and more obscure impressions of the transverse, endocarp fibres, x 1-7. Recent;
Singapore.

FIG. ii. The seed in PI. 10, fig. 6, showing the flat raphe band (r) lying in a shallow
longitudinal furrow of the endosperm. On each side of the band the transverse alignment of
the endosperm is visible. Above where the band has been removed irregular arrangement of
the endosperm is exposed at (ch) and just below longitudinal alignment can be detected.
X 2 approx.

FIG. 12. The same seed, base, showing the aperture for the radicle, the deep furrow (g) due
to the incomplete septum which projected from the locule-wall, and shallow furrows or sinuosities
of the surface. The raphe lay in the furrow at (r). x 1-75.

Anonaspermum aegypticum n. sp.

FIG. 13. The distal end of a seed-cast showing typical ruminate endosperm with encircling
raphe seen at (r, r). The fibrous remains of testa still adhere in the depression at the middle
of the broad surface. The specimen had been fractured transversely and the hilar end was
missing, (ch) indicates the beginning of the chalaza, most of which has been broken away
with the hilar end. x 3. (¥.31106.)

FIG. 14. The same, end view. It shows the encircling raphe partly broken on the left so
that the ruminations are exposed in section at rs. The chalaza begins at (ch). x 3.

FIG. 15. The same, fractured surface, showing the four-partite endosperm, x 3.

Bull. B.M. (N.H.) Geol. 2, 4

PLATE 11

IO

NIPA BURTINI, NIPA FRUTICANS, ANONASPERMUM AEGYPTICUM

PLATE 12

Lagenoidea trilocularis Reid & Chandler

FIG. 16. Side view of a fruit showing three of the six loculicidal segments of the capsule.
X 3. (¥.31107.)

FIG. 17. Base of the same specimen, showing the attachment scar and wrinkled epicarp
which partially obscures the segments. X 3.

Fig. 1 8. Apex of the same, showing more clearly the six segments and lines of dehiscence.

X 3-

FIG. 19. Side view of a second specimen showing three segments clearly ; the edge of a
fourth is just visible on the right-hand margin of the figure. X 2-8. (¥.31108.)

FIG. 20. The same, base, showing six segments and an aperture where placenta and perianth
have broken away. X 2-6.

Lagenoidea bilocularis Reid & Chandler

FIG. 21. A fruit, side, showing the narrow median segment representing the edge of the
septum, and two of the broader lateral segments, x 3. (¥.31109.)

FIG. 22. The same, apex, showing the narrow median segments which form the ends of the
septum and the two pairs of segments which overlie the locules. Loculicidal splitting occurs at
(I, I), septicidal splitting at (s, s). x 3.

FIG. 23. The same, base. The septum is damaged on one side at this end of the fruit.
Lettering as in fig. 22. x 3.

Palaeowetherellia schweinfurthi (Heer) Chandler

FIG. 24. Lower surface of a six-carpelled endocarp showing the six loculicidal suture lines
(/, /). Slight abrasion of this surface has displayed the radiating groups of fine parenchyma
which produce a pitted effect. Decay at the centre of the base has scarcely started. X 2-8.
(¥.31110.)

FIG. 25. The same, apex. The outer part of the carpel wall is corroded in places showing
the finer-celled parenchyma of the layers beneath. Loculicidal sutures are clear ; the septicidal
ones (sp, sp) between them are more obscure except where abrasion has partly removed the
outer layers of the endocarp. x 2-8.

FIG. 26. The under surface of an eight-loculed endocarp, perfect except for the charac-
teristic decay at the centre of the base so that locules and septa (showing planes of weakness
associated with septicidal splitting) are exposed. Several locules have shed their seeds. In
others the seeds are ill-developed, x 2-8. (¥.31111.)

FIG. 27. The base of half a fruit which retains a considerable patch of exocarp (ex) over the
median area. Around the circumference the exocarp has been worn away exposing six seg-
ments of endocarp representing seven locules (/, /). X 2-8. (¥.31112.)

FIG. 28. The fractured surface of the same showing two locules. The one on the right is
occupied by a locule- and seed-cast, (ch) indicates the position of the chalaza. The left-hand
locule is empty but the much encrusted funicle (/) is visible. Its origin in the axis is obscured
by mineral deposit, (ex) = exocarp in section, x 3.

FIG. 29. Base of another incomplete endocarp with three and a half segments representing
four locules (/,/). The half segment on the right has slipped out of position along the loculi-
cidal fracture plane (/). The surface of this half segment shows the plane of septicidal fracture
(sp). X 2-8. (V.3iii3.)

FIG. 30. The same, looking on to the fractured surface, (sp) is the septicidal fracture plane.
(/, /) indicates loculicidal surfaces. The network of fibres is seen on the right-hand loculicidal
suture surface. The left locule surface is obscured by molluscan casts (out of focus in the
figure). X 3.

Bull. B.M. (N.H.) Geol. 2, 4

PLATE 12

LAGENOIDEA TRILOCULARIS, L. BILOCULARIS,
PALAEOWETHERELLIA SCHWEINFURTHI

PLATE 13

Palaeow ether ellia schweinfurthi (Heer) Chandler

FIG. 31. The apex of a nine-carpelled fruit polished by abrasion. (I, 1} indicates planes of
loculicidal dehiscence. Between them, but more obscure, are radial lines indicating planes of
septicidal dehiscence (sp). At (Is) incipient loculicidal splitting can be seen at the middle of
the ridge. The specimen has been fractured along the line (/,/). The small circular scar at
the centre may be the axis or the style base, x 3. (¥.12985.)

FIG. 32. The same, base, corroded at the centre so that the locules are exposed (some now
empty). Note the conspicuous pits over the surface due to the decay of fibres surrounded
by coarse parenchymatous patches. Lettering as in fig. 31. x 3.

FIG. 33. The fractured surface (longitudinal section) of the above showing a locule-cast on
the left with the funicle arising from the axis of the fruit and passing in the plane of the locule
to the hilum situated near the upper outer angle. The right-hand locule is empty, x 3.

FIG. 34. The counterpart half of the same, showing on the right the locule surface which
overlay the cast in fig. 33, and on the left an abortive locule-cast which occupied the empty locule
in fig. 33. x 3.

FIG. 35. The left-hand part of the section in fig. 33. A larger magnification shows the
funicle just above the locule-cast and the entry of the funicle into the cast marking the position
of the hilum on the enclosed seed. This fertile locule lies at (fl) on the left of fig. 32 where the
edge of the cast is seen. Around the hilar end of the cast adherent patches of tissue torn from
the endocarp simulate a rugose testa, x 7.

FIG. 36. The left-hand part of the section in fig. 34. The fibres of the axis show more clearly.
Oblique fine striations on the surface of the abortive locule close to the axis are visible and the
small abortive locule-cast is shown at (a). This locule lies at (fl) on the left of fig. 31. x 7.

FIG. 37. The same as fig. 34, right-hand side of specimen, with a detached seed from
another specimen laid in the locule to show the approximate position that the seed would have
occupied. Axial fibres (a) and funicle (/) are well shown in the endocarp wall and on the flat
surface of the locule respectively. The curved line at the rounded end of the seed is an acci-
dental fracture, x 7. (cf. PI. 14, figs. 43-47.)

FIG. 38. A segment of another fruit which has broken loculicidally, apex. Axis to left at
(a), circumference to right, loculicidal surfaces above and below, x 6. (V. 31114.)

Bull, EM. (N.H.) Geol. 2, 4

PLATE 13

37

32

PALAEOWETHERELLIA SCHWEINFURTHI

PLATE 14

Palaeow ether ellia schweinfurthi (Heer) Chandler

FIG. 39. Side view of the fruit segment in PI. 13, fig. 38. It shows the rounded inner
end of a seed-cast (c) exposed by the breaking away in fossilization of the axis and inner angle
of the carpel. Specimens with similar proportions were separated by Heer (1876) under the
name Royena desertorum. x 6.

FIG. 40. The same, looking on to the inner edge where the two locules and seed-casts can
be seen, x 6.

FIG. 41. Opposite side of the segment to that shown in Fig. 39. Part of the carpel wall
still adheres to the cast at the outer end ; it conceals the hilar end of the seed and embracing
locule-cast, and produces the false effect of a rough nodular testa. The smooth surface at (Ic)
is the remains of the thin locule-cast closely adherent to the seed. To the left at (sc) the film
of locule-cast has broken away, exposing the seed-cast. A narrow band of the smoothly finished
loculicidal surface is preserved at (Is), but elsewhere this surface has been torn away, thereby
exposing the angular cells of the carpel wall at (cw). x 10.

FIG. 42. Apex of another fruit (incomplete as indicated by dotted lines) with exocarp
preserved. Crushing and drying have caused the specimen to crack, white calcite has been
deposited in the cracks which give a false appearance of three perianth segments. The speci-
men had been rubbed down along a plane parallel with its axis at (p). X 1-8. ("^31115.)

FIG. 43. The same, base. A crack on the right exposes a shining seed-cast. This cast is
illustrated laid in the locule of another fruit in PI. 13, fig. 37. Calcite-filled cracks are again
seen. Lettering as in fig. 42. x 1-8.

FIG. 44. Part of the same, more highly magnified and tilted to show the seed-cast of which
the lower margin is exposed. X 6.

FIG. 45. The same seed-cast, removed from the fruit, showing the narrow ventral edge.
The micropyle is at (m), the hilum at (h). The plasticene in which it was necessary to mount
the specimen to obtain this view of the seed somewhat obscures its outline at the lower end of
the figure. X 6-5.

FIG. 46. The same seed-cast, side view, (m) indicates the position of micropyle and radicle,
(h) that of the hilum. The chalaza lay near the lower end of the accidental fracture line. X 10.

FIG. 47. The same cast, opposite surface, to show the bisymmetry of the seed, x 10.

Bull. B.M. (N.H.) Geol. 2, 4

PLATE 14

42

PALAEOWETHERELLIA SCH WEINFURTHI

GEOL. II, 4

15

PLATE 15

Palaeowetherellia schweinfurthi (Heer) Chandler

FIG. 48. The partially polished surface (p) of the fruit in PL 12, figs. 41, 42, showing a
tangential section through a locule and seed. The seed-cast (sc) is closely embraced by the
locule-cast (Ic) lying between the two thick loculicidal valves of the carpel. The raphe (r) is
probably indicated by the slightly constricted area at the narrow upper edge of the seed-cast.
The white mass to the right is calcite filling the loculicidal split and now cementing together
the two valves of the carpel (v, v). x 10.

Icacinicaryayoussefi n. sp.

FIG. 49. A fruit, broad surface, showing form and ornamentation, (st] indicates the posi-
tion of the style. The funicle was situated in the right margin of the enclosed endocarp. x 2-6.
(V.3ui6.)

FIG. 50. The same, opposite surface. Contraction of the pericarp has brought out the
outline of the endocarp within, x 2-6.

FIG. 51. An endocarp bereft of the pericarp. Note the surface ornamentation, the basal
aperture for the entry of the funicle at (/), and the short vertical fracture with infiltrated
mineral substance at the sty lar end (st). x 2-6. (¥.31117).

Icacinicarya sp. ?

FIG. 52. Side view of a wrinkled drupe showing the asymmetry characteristic of Icacinaceae.
The more convex left margin may be presumed to carry the funicle. (st) indicates the style.
X 2-8. (V.3in8.)

FIG. 53. Base of the same, showing the inflated funicular margin (below) and narrow
opposite margin (at the top of the figure). The small circular scar of attachment, from which
the cells radiate, is seen at the centre, x 3.

FIG. 54. Apex of the same. Funicle bearing margin to the left. Style at (st). x 2-6.

Carpolithus sp.

FIG. 55. Fruit, side, showing an angle at the apical end and small angular concavities all
over the surface, x 2-8. (¥.31119.)

FIG. 56. The same, apex, showing the three angles or ridges and the surface concavities.
X 2-6.

FIG. 57. The same, base, showing the slightly sunk attachment and the surface concavities,
as well as the shallow grooves which lie opposite the apical angles, x 3.

Bull. B.M. (N.H.) Geol. 2, 4

PLATE 15

50

49

PALAEOWETHERELLIA SCHWEINFURTHI, ICACINICARYA YOUSSEFI,
ICACINICARYA SP. ? CARPOLITHUS SP.

PLATE 16

Thiebaudia rayaniensis n. gen. et sp.

FIG. 58. Apex of fruit. The pericarp is preserved only at the centre. It shows shallow,
inconspicuous, radial furrows diverging from a slightly sunk circular area (perhaps the style
base). Where the pericarp is broken away, the pulpy mass which fills the fruit cavity is exposed.
On the surface of this mass radial furrows mark the position of shallow longitudinal ridges which
must have projected from the fruit-wall. Between them impressions of the placentae are seen
each with two rows of seeds now represented by the hollows which they produced on the surface
of the pulp. Each placenta arises from a short tongue of tissue with longitudinal median
furrow. These tongues project from beneath the edge of the remaining pericarp, x 1-6.
(V.3H20.)

FIG. 59. The same, from below. Here the pericarp is preserved only at the very centre of the
base. The radial furrows (so clear on the upper surface) are obscure, for this surface has been
much battered and partially destroyed. The arrangement of the seeds has been disturbed, but
a few external seed impressions and internal casts are preserved scattered somewhat irregularly.
A small concavity, seen at the lower edge of the central fragment of the pericarp, may be the
external impression of a detached seed (cf. fig. 62). x 1-6.

FIG. 60. Part of the upper surface (indicated by (x) in fig. 58). It shows more clearly the
seed pockets on each side of the placentas, and the impressions of the ridges described above.
The margin of the fragment of pericarp lies near the base of the figure at (p). x 6.

FIG. 61. Part of the under surface (indicated by (x) in fig. 59) showing a few of the hollow
pockets for the seeds. They sometimes show a central projection (best seen in the hollow
marked s) which suggests a concavity on the surface of the seed. A few broken fibre-fragments
(remains of pericarp) adhere to the pulpy mass, x 6.

FIG. 62. The external impression of the seed (?) on the pericarp in fig. 59 as described
above. The surface of the pericarp was painted white around the impression to bring out its
outline. A central depression must correspond with a prominence on the seed itself. Coarse
cells or areoles are obscurely seen diverging from this central depression, x 16.

FIG. 63. A somewhat imperfect internal cast of a seed from the lower surface of the pulpy
mass. Striae due to cells on the surface of the cast lie parallel with its longer axis. Although
the cast is obscure a sharp marginal angle suggests a bisymmetric form, x 20. (V.3U2O.)

Carpolithus hassani n. sp.

FIG. 64. The exterior of a valve showing three deep furrows and a branching fibre which
is partially embedded in the surface between the right-hand furrow and the short middle one.
X 1-8. (¥.31121.)

FIG. 65. The internal aspect of the same, showing that the wall is pierced by the three
external furrows. The thickness of the wall is also shown. The smooth marginal areas appear
to have been artificially produced by polishing, x 1-8.

Carpolithus sp. (Icacinicarya sp. ?)

FIG. 66. The internal cast of a valve of a fruit showing form and ornamentation suggestive
of Icacinaceae. x 1-8. (¥.31122.)

Bull. EM. (N.H.) Geol. 2, 4

PLATE 16

THIEBAUDIA RAYANIENSIS, CARPOLITHUS HASSANI
CARPOLITHUS SP. (ICACINICARYA SP. ? )

.

PRESETED
1 6 MAR 1954

PRINTED IN GREAT BRITAIN BY
ADLARD AND SON. LIMITED
BARTHOLOMEW PRESS, DORKING

5 OCi 1954

THE CARBONIFEROUS
FLORA OF PERU

W. J. JONGMANS

BULLETIN OF
THE BRITISH MUSEUM (NATURAL HISTORY)

GEOLOGY Vol. 2 No. 5

LONDON: 1954

BULLETIN OF
THE BRITISH MUSEUM (NATURAL HISTORY)

GEOLOGY

The following papers appeared in Volume I (1949-52) :

Price
No. I (1949). The Pterobranch Rhabdopleura in the English Eocene.

H. D. Thomas & A. G. Davis 75. 6d.

No. 2 (1949). A Reconsideration of the Galley Hill Skeleton. K. P.

Oakley & M. F. Ashley Montagu 5$-

No. 3 (1950). The Vertebrate Faunas of the Lower Old Red Sandstone

of the Welsh Borders. E. I. White.

Pteraspis leathensis White a Dittonian Zone-Fossil. E. I.

White • 75- 6d'

No. 4 (1950). A New Tithonian Ammonoid Fauna from Kurdistan,

Northern Iraq. L. F. Spath .... . IDS.

No. 5 (1951)- Cretaceous and Eocene Peduncles of the Cirripede Euscal-

pellum. T. H. Withers .55.

No. 6 (1951). Some Jurassic and Cretaceous Crabs (Prosoponidae).

T. H. Withers ... 5*-

No. 7 (1952). A New Trochiliscus (Charophyta) from the Downtonian

of Podolia. W. N. Croft . . IDS.

No. 8 (1952). Cretaceous and Tertiary Foraminifera from the Middle

East. T. F. Grimsdale . IDS.

No. 9 (1952). Australian Arthrodires. E. I. White . . . 155.

No. 10 (1952). Cyclopygid Trilobites from Girvan. W. F. Whittard . 65.

THE CARBONIFEROUS FLORA OF PERU

BY

WILHELMUS JOSEPHUS JONGMANS

Pp. 189-224 ; Pis. 17-26

BULLETIN OF
THE BRITISH MUSEUM (NATURAL HISTORY)

GEOLOGY Vol. 2 No. 5

LONDON : 1954

THE BULLETIN OF THE BRITISH MUSEUM
(NATURAL HISTORY), instituted in 1949, is
issued in five series corresponding to the Departments
of the Museum, and an Historical Series.

Parts will appear at irregular intervals as they become
ready. Volumes will contain about three or four
hundred pages, and will not necessarily be completed
within one calendar year.

This paper is Vol. 2, No. 5 of the Geological series.

PRINTED BY ORDER OF THE TRUSTEES OF
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Issued September 1954 Price Fifteen Shillings

THE CARBONIFEROUS FLORA OF PERU

By W. J. JONGMANS

SYNOPSIS

A reconsideration of all the older records of Carboniferous plants from Peru, together with
an examination of two important newer collections, fully confirms the Lower Carboniferous
age of the flora. Several new species are described, including representatives of Lepidoden-
dropsis, and the world distribution of this genus is reviewed.

INTRODUCTION

THE Carboniferous flora of Peru has been the subject of several papers. The principal
flora is found on the peninsula of Paracas, which, as Berry (1922) points out, is
largely made up of continental Carboniferous sediments and constitutes one of the
very few deposits of this character in South America, and the only known occurrence
of rocks of this age on the west coast of South America. Somewhat to the north-east
of the peninsula there are other Carboniferous localities at Huanuco and in the
neighbourhood of Cerro de Pasco. Up to the present little has been known of the
flora of these localities. Gothan (1928) mentions Rhacopteris circularis Walton
from Vichaicoto and S. Huanuco and Knorria and a Calamites-like specimen from
Cachama, between Cerro de Pasco and Huanuco. Fortunately one of my former
assistants, Dr. N. de Voogd, sent me a good collection from Carhuamayo. This
collection and another from Paracas, put at my disposal by the Trustees of the
British Museum, will be described in this paper.

REVIEW OF THE LITERATURE ON THE CARBONIFEROUS FLORA

OF PERU

Almost at the same time as Berry, Seward (1922) described another set of plants
from Paracas. According to these papers the outcrop at Paracas was first discovered
by Fuchs (1900). Seward states : "The coal occurs in a series of greenish sandstones
and grey and black carbonaceous shales, which have a north-easterly strike and dip
about 25° south-eastwards. These are overlain unconformably on the neck of the
peninsula by Tertiary sandstones and impure limestones. There is no definite
stratigraphical evidence of the age of the coal-bearing beds, and the plants are there-
fore of special importance."

Fuchs (1900) recorded the following species : Catamites suckowi Bgt., Sphenopteris
hartlebeni Dkr., Baiera pluripartita Schl., Lepidodendron sternbergii Bgt., Sigillaria
tessellata Bgt., and Stigmaria ficoides Bgt.

GEOL. II, 5. j6

IQ2 THE CARBONIFEROUS FLORA OF PERU

Fuchs assigned the beds to the Upper Coal Measures, but his list is scarcely pos-
sible ; it contains common Carboniferous plants and Wealden species.

The Carboniferous flora was mentioned subsequently by Fuchs (1905), Dorca
(1909), Marsters (1909) and Lisson (1917), without any reference to the Wealden
species. Lisson's paper is the only one which contains any new facts on this flora ;
he mentions Lepidodendron rimosum and L. obovatum, determined by Zeiller. The
age is given as Westphalian. Some critical remarks will be given when treating
the papers by Berry (1922) and Gothan (1928).

The next contribution to the Carboniferous flora of Peru was that by Steinmann
(1911) who did not visit Paracas. He examined Fuchs' collection at the Cuerpo
de Ingenieros de Minas in Lima, and listed the following forms for the Paracas
flora : Archaeocalamites radiatus, Lepidodendron cf. veltheimi, L. cf. volkmanni,
Sphenopteris affinis (furcata}, Rhodea filifera and Rhabdocarpus.

The occurrence of these species would prove that the flora belongs to the Lower
Carboniferous. It is interesting but somewhat remarkable that the plant-bearing
Carboniferous should be of Lower Carboniferous age and the invertebrate-bearing
Carboniferous of Upper Carboniferous age. Unfortunately no literature on the
invertebrate fauna of the Peruvian Carboniferous is known to me, and so far as can
be seen from the sections in Berry's papers (1922, 1922^) no invertebrate fauna
had been met with on the peninsula of Paracas during his visit. Steinmann states
that no marine fossils are known from Paracas. He visited a second Carboniferous
locality near Huichaycota, some kilometres south of Huanuco on the Huallaga and
found large stems of Lepidodendron and at different places many specimens which
he records as Rhacopteris inaequilatera Goepp.

The introductory paper by Berry (i 922*1) contains no descriptions of the plants,
but discusses the locality and the history of the knowledge of the flora up to that
date and records the following :

Palmatopteris furcata (Bgt.), Eremopteris whitei Berry, E. peruianus Berry, Cata-
mites suckowi Bgt., Calamostachys sp., Lepidodendron rimosum Sternb., L. obovatum
Sternb., Lepidophyllum sp., Lepidostrobus sp., Stigmaria sp. and Knorria sp.

Berry visited Paracas and was able to make large collections ; his remarks on the
locality and the mode of occurrence of the plants are very valuable. According to
Berry the fossil plants occur at different horizons in the sections he examined, and
" there is no chronologic change in the flora from top to bottom although fossil
plants are more varied in the lowermost horizon. The materials are relatively
coarse throughout and would seem to indicate rapid deposition." Over half of the
section " is described as sandstone, which is often coarse and arkosic. Of the 273
feet described as shale 192 feet are distinctly sandy, so that less than 14% of the
total thickness [of 585 feet], including the so-called coal seams, is fine-grained shale
and even the coal contains much silty impurities. No underclays with rootlets, or
upright stems were observed and the coaly layers have every appearance of having
been formed of drift material " (Berry, 1922^ : 191-192).

The flora is " extremely limited, although some of the elements are exceedingly
common," especially the plants recorded as Palmatopteris furcata, Eremopteris
whitei, Catamites suckowi and Lepidodendron rimosum (Berry, 1922^ : 193). Berry

THE CARBONIFEROUS FLORA OF PERU 193

imagines " that the coarseness of the sediments and the apparent drifting of the
material are mainly responsible for the absence of a more representative flora."
Thus his collections contained " no traces of Sigillaria, Cordaites, Sphenophyllum,
Catamite foliage, nor of any Neuropterids, Pecopterids, Alethopterids or Lonchop-
terids." He states that this feature of the flora is undoubtedly responsible for
Steinmann's opinion (1911) that the Paracas flora is of Lower Carboniferous age.
According to Berry the Paracas Carboniferous corresponds to the Westphalian stage.

In the same year Seward (1922) described a collection of plants from Paracas made
by J. A. Douglas in 1911. Seward's flora contains Sphenopteris sp., Lepidodendron
sp., Sigillaria (or Lepidodendron) sp., Bothrodendron sp., and Planta incertae sedis.
His determinations are considered in detail below :

Sphenopteris sp. (Seward, 1922, pi. 13, figs. 1-3) is present in several fragments.
" The branched axis is longitudinally striated and smooth ; the pinnules are more
or less deltoid, deeply dissected, and the ultimate segments are obtuse or truncate."
Seward compares it with Sphenopteris furcata Bgt., but in this species the segments
are acute. He also compares it with Eremopteris missouriensis Lesquereux (White,
1899, pi. 5, figs. 1-30) which does possess obtuse or truncate ultimate segments.

Several Lower Carboniferous species have pinnules which closely resemble the
Peruvian specimens ; " the deeply dissected form of the lamina suggests comparison
with pinnules of Rhodea and Sphenopteridium." Thus Nathorst (1920, pi. i, figs,
11-13) figures as Sphenopteridium norbergii a plant which he compares with Sphen-
opteris affinis L. & H. ; the latter differs in its broader, thicker segments, and its
stronger and more numerous nerves. It will later be shown that such comparison
is important for the determination of the specimens and their age.

Lepidodendron sp. (Seward, 1922, pi. 13, figs. 4-6). The most important specimen
(fig. 4) shows a branch with attached leaves. It is preserved on a carbonaceous
sandstone and most of the details cannot be seen. However, it is clear that the
" cushions " of the leaves are separated by undulate lines, that succeeding " cushions"
are connected at the upper and lower ends, and that the surface is striated, though
not in the deeply impressed portions of the cushions. Seward's fig. 5 shows some
details on the leaf-cushion and the leaf-scar.

On pi. 13, fig. 6 Seward figures a Lepidodendron which he considers belongs to
the same species as those illustrated in figs. 4, 5. The specimen is interesting as it
shows the leaf-like organs very clearly. It somewhat resembles the figure published
by Johnson (1913, pi. 41, fig. 3), but it is impossible to decide whether it is a portion
of a cone or not. The long erect leaves are narrow and possess a distinct middle
nerve. They are sharply pointed at their ends.

Sigillaria (or Lepidodendron) sp. (Seward, 1922, pi. 13, figs. 7, 8). " Pieces of
a stem having contiguous leaf-cushions which bear leaf-scars agreeing both with
some types of Sigillaria (e.g., S. brardi Bgt.) and with certain species of Lepidoden-
dron. On the upper part of several leaf -cushions there is a small circular scar,
presumably a ligule-pit. No vascular bundle scars or parichnos-scars can be detec-
ted." Seward compares his specimens with those named 5. brardi from South
Africa (Seward, 1897 : 326) and Brazil (White, 1908), which, however, probably
do not belong to Brnogniart's species, nor even to the genus Sigillaria. He also com-

194 THE CARBONIFEROUS FLORA OF PERU

pares them with S. mutans Weiss & Sterzel (1893 : 84). It is very curious that
Seward should compare his specimens with a true Stephanian species.

Bothrodendron (?) sp. (Seward, 1922, pi. 13, fig. 9 ; text-fig.). Seward describes
two specimens under this heading " although it is by no means certain that the
specimens shown in pi. xiii, fig. 9, and in the text-figure belong to the same species."

The specimen figured in pi. 13 shows " spirally-disposed and widely-separated,
slightly prominent, transversely elongated, rhomboidal leaf-scars. " On the right-
hand side, a thin carbonaceous layer probably represents the actual surface . . .
On the partly decorticated surface there are discontinuous longitudinal ridges,
and an irregular transverse wrinkling, but on the carbonized film no wrinkling is
seen. There is no indication of any leaf-cushion, no ligular pit, and only a very faint
suggestion in a few of the scars of a median vascular scar. The leaf-scars shown
in the text-figure are rather more rounded and appear as slightly concave areas . . .
In the small and widely-separated leaf-scars these fragments agree with Both-
rodendron, Pinakodendron, and Asolanus. The form of the leaf -scar and the absence
of a leaf-cushion are features more suggestive of Bothrodendron."

The features mentioned by Seward are those which are shown on the stems and
larger branches of Bothrodendron, whereas clear leaf-cushions and leaf-scars of a
more lepidodendroid form occur on the smaller branches. It is known that the gap
between these two extreme forms is filled by a whole series of transitions. It is
somewhat tempting to consider the three forms described by Seward as smaller
branches and stems of the same species.

The last plant mentioned by Seward (1922, pi. 13, fig. 10) has not been named.
This is represented by crowded branched filaments which are " probably portions of
pinnules of a fern-like plant, such as some of the Lower Carboniferous species referred
to Rhodea or Sphenopteridium."

Seward was at first inclined to regard his material as Upper Carboniferous in age.
Dr. Kidston, however, regarded the palaeobotanical evidence as more favourable
to a Lower Carboniferous horizon. After a re-examination of the specimens Seward
modified his first opinion and agreed with Kidston. He remarks, however, that
the plants are too imperfect to serve as trustworthy guides and that " further
research is greatly to be desired, since the available data are inadequate as a basis
for any positive statement."

Shortly after Seward's paper appeared, a fully illustrated account of the flora
was published by Berry (1922). The determinations are discussed below :

Palmatopteris furcata (Bgt.) Berry, 1922, pi. i, figs. 1-3.

The plant he named Palmatopteris furcata is exceedingly common in the Paracas
deposits. He unites with it Sphenopteris affinis (Steinmann, 1911 : 50) and Sphen-
opteris hartlebeni (Fuchs, 1900 : 50). These records certainly represent the same
species, but it is not certain that the specific determination given by Berry is right.
He does not give a detailed description and an opinion can be based only on his
figures (pi. i, figs. 1-3). The species has since been renamed Sphenopteris paracasica
by Gothan (1928 : 293).

THE CARBONIFEROUS FLORA OF PERU 195

Eremopteris peruianus and E. whitei (Berry, 1922, pis. 2-4).

Berry describes two new species of Eremopteris. He points out the difficulties
of demarcation between Eremopteris (especially as used by American authors) and
Rhacopteris. He compares his Eremopteris peruianus (pis. 2, 3) with Rhacopteris
transitions (Ett.) as recorded by Stur (1875, pi. 8, figs. 5-7) but there are other
species of " Anisopteris " with which Berry's material may be compared. Accord-
ing to his description and figures the size and form of the pinnules are very variable
and it is quite likely that more than one species of Rhacopteris occurs in the material.

According to Berry his second species, Eremopteris whitei, is identical with E.
elegans Lesquereux (1880, pi. 53, fig. 7). Berry considers this species to be entirely
different from the European Rhacopteris elegans (Ett.) and from R. (Sphenopteris}
asplenites Gutb. These European species have nothing in common with the Peru-
vian material.

Berry describes his material as follows : " The pinnae are linear oblong, their
divisions or pinnules are oblique, oblong or rhomboidal in form, narrowed to the
somewhat decurrent base, deeply pinnately cut by narrow sinuses into cuneate
divisions which are rounded or subcrenate distad. The venation is flabellate and
largely immersed in the thick substance of the lamina."

Eremopteris whitei Berry cannot be a Rhacopteris (sensu stricto) or Anisopteris.
It might possibly be compared with some species of Sphenopteridium as figured by
different authors (e.g., Walton, 1926) but Berry's figure alone is not sufficient to
identify it with Eremopteris or Rhacopteris, and until further data are available it
can be named Sphenopteris whitei (Berry).

It must have been a rather robust plant as evidenced by the stout longitudinally
striated rachises. The nervation, as far as can be seen in Berry's figure, does not
seem to agree with that of Rhacopteris or Anisopteris nor with Eremopteris elegans
as figured by Lesquereux (1880, pi. 53, fig. 7) ; it can much better be compared with
that of Sphenopteridium. Another plant with which it may be compared is that figured
by me (1940, pi. 4, fig. 9) from the Lower Carboniferous of Egypt and erroneously
referred to Rhodea cf. hochstetteri Stur. Better material which I received later
shows that the Egyptian specimens cannot belong to Stur's species, and I now have
little doubt as to their identity with Berry's species.

Calamites suckowi Bgt. (Berry, 1922, pis. 5-7).

The plants figured by Berry as Calamites suckowi Bgt. are very fragmentary and
broken. Most of them do not show a nodal line. The best example is seen in pi. 6.
Here a nodal line is present on which the sharply pointed ribs are clearly alternating,
so that a reference to Asterocalamites is precluded. The stems undoubtedly belong
to Calamites, but the sharply pointed ribs show that it cannot be C. suckowi ; it
looks more like C. undulatus Sternb.

Lepidodendron rimosum Sternb. and L. obovatum Bgt. (Berry, 1922, pi. 8 ; pi. i,

fig- 5).

The two species of Lepidodendron, L. rimosum Sternb. (plate 8) and L. obovatum
Bgt. (pi. i, fig. 5) described and figured by Berry agree with Zeiller's opinion on this

ig6 THE CARBONIFEROUS FLORA OF PERU

material (1917). It must be pointed out, however, that if Berry's figures agree
with the originals there is a remarkable difference between pi. i, fig. 5, and pi. 8,
figs, i, 2, on the one hand and pi. 8, fig. 3, on the other. Only in pi. 8, fig. 3, is a leaf-
scar visible and it is likely that this drawing was not made from the figured specimens
but included only to indicate Berry's conception of the species. His determinations
will be discussed after reference to Gothan's paper.

Lepidostrobus sp. (Berry, 1922, pi. i, fig. 4).

In his pi. i, fig. 4, Berry figures a Lepidostrobus, described as " A characteristic
Lepidodendron cone, too poor for identification or description." It is by no means
certain that it is a Lepidostrobus. The cone is very similar to one figured by Johnson
(1913, pi. 41, figs. 3, 4) as a cone or strobilus of Bothrodendron kiltorkense, but it
does not show sufficient details to be certain of its identity.

Gothan (1928) described and figured some Carboniferous plants collected by
Steinmann from Paracas and other localities to the north of this peninsula. Stein-
mann (1929) also dealt with this material. As many of the latter's figures are much
clearer than those in Gothan's paper, the two can be considered together. A short
review of the descriptions and discussions follows here :

Rhodea sp.

This is the form already recorded as Rhodea filif era by Steinmann (1911) and as
Planta incertae sedis by Seward (1922). It is quite possibly a Rhodea, but specific
determination is not possible.

Sphenopteris paracasica Gothan (1928, pi. 13, fig. i) (including Palmatopteris
furcata Berry, 1922, pi. i, fig. i ; Sphenopteris sp. Seward, 1922, pi. i, figs, i, 2 ;
Steinmann, 1929, text-fig. 30^, b}.

Steinmann's figure is a copy of that in Gothan's paper, together with an enlarge-
ment of the pinnules. Seward, who notes the difference from Palmatopteris furcata,
compares his material with Rhodea or Sphenopteridium, such as 5. norbergii Nath.,
a species which its author compared with Sphenopteris affinis. Gothan com-
pares it with Adiantites bellidulus Hr. (Nathorst, 1914, pi. 15, figs. 11-14). It is
very probable that Sphenopteris affinis as recorded by Fuchs (1900) belongs to
Gothan's species. Another species with which Gothan compares it is Sphenopteris
bodenbenderi Kurtz (1921) from Carrizal, near Famatina, La Rioja. A comparison
is also possible with some of the species described by Walton (1931, especially pi.
25, fig. 23) from the British Lower Carboniferous. The specimens known so far are
too fragmentary for determination.

Nevertheless there is an important difference in the rachis as figured by Gothan
and that figured by Seward. The former shows a rachis with a distinct middle line
and no other ornamentation. Seward shows a rachis with a longitudinal striation.
It is by no means certain that these two sets of specimens and those described by

THE CARBONIFEROUS FLORA OF PERU 197

Berry (no rachis and more penicillate pinnules) belong to the same species. There
may be three different species, and so for the present it would be best to use Gothan's
name Sphenopteris paracasica to indicate this type of plant.

Rhacopteris circularis Walton (Gothan, 1928, pi. 15, fig. i).

Recorded from Vichaicoto, S. Huanuco, but not from Paracas, this is without
doubt a Rhacopteris, though Gothan's specific determination may be questioned.
Steinmann's fig. 29 includes a specimen which is comparable with that of Gothan's,
together with some other figures of details of the leaves ; organs which he considers
as aphlebia (D and E) ; and a fragment of a rachis (F), all found together with
R. circularis. These figures, however, show details which are not completely
identical with those in Gothan's figure.

The occurrence of Rhacopteris in these floras is very important. Berry (1922,
pi. 2, figs. 1-3) described Eremopteris peruianus from Paracas, which almost certainly
belongs to Rhacopteris and should be named Rhacopteris peruianus (Berry). It is
possible, however, that it contains more than one species. The form of the pinnules
is very variable, and as Gothan (1928 : 297) points out, it is probable that Berry's
pi. 2, figs. 2, 3, especially fig. 3, belong to the plant which Gothan named Rhacopteris
circularis Walton. If this is true the species has also been found in Paracas.

Berry's pi. 2, fig. i can be compared with Rhacopteris petiolata (Walton, 1926,
pi. 16, fig. 10). For the present, however, Berry's name Rhacopteris peruianus
must be retained, though new material may eventually show that it belongs to some
previously described species.

Calamites peruvianus Gothan (1928, pi. 14, fig. i).

Gothan illustrates some poorly preserved fragments of Calamites as a new species
Calamites peruvianus, and considers Berry's C. suckowi as identical. As already
pointed out (p. 195) these fragments belong to Calamites and not to Mesocalamites
or Asterocalamites, since the ribs alternate regularly at the nodal lines. Their ends
are not blunt as in C. suckowi, and Berry's figures show the sharp pointed ends of
the ribs as in C. undulatus. Gothan supposes that these Calamites are only distantly
related to the European species. However, the characters on which he bases this
opinion (thin stems, with long and narrow internodes, without branch-scars or infra-
nodal canal-traces) are characters which can be found in C. undulatus or which may
be due to poor preservation and the fragmentary state of the specimens. There
is no reason to create a new specific name for these fragments or to compare them
with Phyllotheca or other genera of Equisetales. The specimen figured by Steinmann
(1929, text-fig. 28) is even more fragmentary than those mentioned above. The
only name which might be given is Calamites cf. undulatus Sternb., though the correct
name should be Calamites sp. indet.

Lepidodendron peruvianum Gothan (1928, pi. 13, fig. 2).

This specimen is interesting as it does not show a leaf-scar. Gothan unites with
this species L. obovatum and L. rimosum of Berry (1922, pi. i, fig. 5 ; pi. 8, fig. i

ig8 THE CARBONIFEROUS FLORA OF PERU

only). Berry's pi. 8, fig. 3 does not belong to this species, and most probably the
drawing has nothing to do with the specimens figured on the same plate. In my
opinion, however, Berry's pi. 8, fig. 2 should also be united with Gothan's species.

No leaf-scar is visible and the broad bands between the leaf-cushions point to
Lepidodendropsis. The specimens may be compared with Lepidodendropsis fenestrata
from the Lower Carboniferous of Egypt (Jongmans & Koopmans, 1940, pi. 2, figs.
4#-c). The only real difference is that those described by Berry and by Gothan
are larger, and the leaf-cushions are broader and not so elongated. Gothan states
that the surface of the bands which separate the cushions is smooth.

The figures published by Steinmann (1929, text-figs. 23A-D) are much better ;
they show clearly that the material belongs to Lepidodendropsis and the similarity
to the Egyptian material is still more striking. The Peruvian species should
therefore be named Lepidodendropsis peruviana (Gothan) Jongmans.

Lepidodendron lissoni Steinmann (Gothan, 1928, pi. 14, fig. 2).

Another species of Lepidodendron figured by Gothan (1928) and by Steinmann
(1929, text-fig. 24) is L. lissoni Steinmann. Gothan compares it with L. spetsber-
gense Nath. So far as one can judge from the figure the specimen is badly preserved
and it may be an older stem of Bothrodendron.

Lepidodendron sp. (Gothan, 1928, pi. 15, fig. 3, left).

It is not possible to give an opinion on this specimen. There may be some con-
nexion with Seward's pi. 13, figs. 7, 8 (Sigillaria or Lepidodendron), but this will be
very difficult to prove. Gothan also refers to a branch of Lepidodendron which
he compares with Seward's pi. 13, figs. 4, 6. He does not figure the specimen.

Asolanus (?) minimus Gothan (1928, pi. 15, figs. 2, 20,, ? 3, right).

The material is insufficient for a true understanding of the plant which Gothan
named Asolanus (?) minimus ; it is not clear that it is an Asolanus. The ornamenta-
tion is more like that of Bothrodendron (or Cyclostigma} than of Asolanus.

? Bothrodendron pacificum Steinmann (Gothan, 1928, pi. 13, figs. 3, 3#).

Gothan compares his specimen with Bothrodendron sp. (Seward, 1922, pi. 13,
fig. 9). It is very probable that Seward's figure represents a Bothrodendron- or
Cyclostigma-like stem, but since all the leaf-scars in Gothan's figure are broken it is
not possible to identify details of the scar. A ligule-scar could not be observed.

Fortunately Steinmann (1929, text-fig. 27) gives a much better picture of "Bothro-
dendron " pacificum. His figure is similar to Seward's fig. 4 and must belong to
the same species. It follows that Steinmann's specific name pacificum can be used
for Seward's (pi. 13, figs. 4, 7, 8), Gothan's (pi. 13, figs. 3, 3^) and Steinmann's
figures. With the description of new material to be dealt with later in this paper,
it will be shown that this species must be put into Cyclostigma and that Seward's
fig. 9 is a bigger stem belonging to the same species. There is no trace of a ligule
and therefore the material must belong to the eligulate group of the Lycopodiales.

THE CARBONIFEROUS FLORA OF PERU 199

Trachyphyton neglegibile Gothan (1928, pi. 14, figs. 3, 4).

It is difficult to give an opinion on Gothan's Trachyphyton neglegibile. He compares
it with Stigmariopsis. No root-scars are visible, though these are often very
indistinct in specimens of Stigmariopsis. Johnson (1914, pi. 15) figures some
rhizomes of Cyclostigma kiltorkense which closely resemble Gothan's figure.

Gothan considers the flora to be of Lower Carboniferous age.

A further contribution to the Carboniferous flora of Paracas was published by
Read (1938). It is based on material collected from the classic locality by H. Bassler
in 1922. Bassler did not make a special exploration but picked only debris of rocks
from a shallow shaft made by former collectors. Read records the following plants :
Sphenopteris " parasica" Gothan ( = S. paracasica), Adiantites whitei (Berry),
Adiantites peruianus (Berry), Adiantites bassleri Read, Rhacopteris ovata (McCoy)
Walkom, Rhacopteris cf. cuneata Walkom, Aphlebia australis Read, Lepidodendron
peruvianum Gothan and Catamites peruvianus Gothan.

The type of flora described by Read is much the same as that in earlier publica-
tions. Some of his determinations are discussed below.

Adiantites whitei (Berry) and A. peruianus (Berry) Read, 1938, text-fig. 3.

It is not entirely clear why Read includes Eremopteris whitei and E. peruianus
in Adiantites. In my opinion it would be better to unite E. peruianus with Rhaco-
pteris. I am not sure that Read's fig. 3 is the same plant as Berry's E. peruianus
and I am inclined to compare it with Triphyllopteris collombiana Sch. As to Eremo-
pteris whitei I prefer to leave this in Sphenopteris until better and more complete
specimens have been found.

Adiantites bassleri Read (1938, text-fig. 7).

Adiantites bassleri Read is known only from a few fragments. They show the
outline but no venation. This plant possibly belongs to the species which will be
described as Triphyllopteris collombiana (Sch.) (p. 214).

Rhacopteris ovata (McCoy) Walkom (Read, 1938, text-fig, i).

I entirely agree with Read's determination of his text-fig, i as Rhacopteris ovata
(McCoy). As will be seen later, new material from the De Voogd collection clearly
shows that this form is represented in the flora. The specimen figured by Gothan
(1928, pi. 15, fig. i) as R. circularis Walton does not show the same characters but
they are seen in Steinmann's text-fig. 29, in Berry's pi. 2, fig. 3 (? 2), and in the speci-
mens figured in this paper.

It is possible that Gothan's figure should be separated from the others and com-
pared with R. circularis Walton, or it may be that the margins in Gothan's specimen
are not well preserved.

It is curious that Read states that Rhacopteris ovata is very common in Paracas.
So far as is known from the older literature there is only one figure from Paracas
(Berry's E. peruianus, pi. 2, fig. 3 ; fig. 2 is very doubtful) which may be compared

200 THE CARBONIFEROUS FLORA OF PERU

with R. ovata. Steinmann's specimens and Gothan's R. circularis Walton are from
Vichaicoto, S. Huanuco.

Rhacopteris sp. cf. JR. cuneata Walkom (Read, 1938, text-fig. 5).

A single fragment figured by Read as Rhacopteris sp. cf . R. cuneata may be correctly
determined. Two fragments which resemble Read's figure will be described with the
new material from Paracas belonging to the British Museum.

Aphlebia australis Read (1938, text-fig. 6).

This is a curious plant. It is a leafy, highly divided fragmentary organ ; the
base is unknown. It is not possible to show a relationship with the other elements
of the flora.

Lepidodendron peruvianum Gothan (Read, 1938, text-fig. 4).

Read's figure is not exactly like Gothan's pi. 13, fig. 2, and is more like Steinmann's
figures. It is probable that this specimen belongs to Lepidodendropsis.

Catamites peruvianus Gothan.

Read mentions some specimens of Calamites which he unites with C. peruvianus
Gothan. As already stated a specific name is unnecessary for these fragments.

In a paper on the fossil plants of the Neo-Paleozoic of Brazil, Read (1941 : 17)
reviews the Mississippian floras of South America and gives a list of the Paracas
flora according to his own observations. He records a similar flora collected from
10 km. N.E. of Garhuamayo, Peru. The collection is fragmentary and the following
species are noted : Rhacopteris ovata (McCoy), Adiantites bassleri Read and Lepi-
dodendron peruvianum Gothan.

He compares the flora with Lower Carboniferous floras from Argentina. Such
a comparison could be made with success only after a revision of the floras of that
country, since so many records from Argentina are in need of correction. It is
clear, however, that the lower portion of the Carboniferous in Argentina has a similar
Rhacopteris-ftora., probably also with Lepidodendropsis to which almost certainly
some of the Lepidodendron species listed from there may prove to belong.

Frenguelli (1943) was able to study some specimens of Rhacopteris from Carhua-
mayo, Peru, in the Museum of La Plata. (From this locality I received the speci-
mens which are described in the last section of this paper). Frenguelli figures two
specimens ; his pi. 3, fig. i, is named Rhacopteris circularis Walton according to
Gothan's determination and figure (1928, pi. 15, fig. i), and his pi. 3, fig. 2 is named
R. ovata and agrees with Steinmann's figures (1929, text-figs. 29A-c) and with Berry's
pi. 2, fig. 3, and perhaps fig. 2 (1922). It is therefore probable that both species
occur in the Lower Carboniferous of Peru.

In the same paper Frenguelli discusses specimens from Argentina. The specimen
figured in his pi. i and the poorly preserved fragment in pi. 2, fig. i, are from the
Mine El Tupe, La Rioja, and those in pi. 4, figs. 1-3, are from Agua Salada, La
Rioja ; these he names Rhacopteris ovata (McCoy). On pi. 4, fig. 4, he figures a

THE CARBONIFEROUS FLORA OF PERU 201

specimen also from Agua Salada which he names R. circularis Walton. The differ-
ence between the two species is rather slight. It may be that the crenulate border
of R. ovata is different from the more or less entire margin of R. circularis and that
the form of the segments is somewhat different from those of R. circularis which
are more circular. At all events it is interesting that the Argentine species of
Rhacopteris agree with the Peruvian ones.

Frenguelli (1943, pi. 2, fig. 3 ; pi. 4, fig. 4) also figures specimens which he calls
Calamites peruvianus Gothan, but they are equally as indeterminable as all or at
least most of the other figures published under this name. It is useless to name or
figure such bad specimens.

In an earlier paper Frenguelli (1941) also compares Peruvian plants with the
flora of Argentina, describing an old collection of fossil plants from Agua de los
Jejenes, San Juan. This flora is said to contain : Rhacopteris semicircularis Lutz,
R. ovata (McCoy), Eremopteris cf. sanjuanina Kurtz, E. cf. whitei Berry, Rhabdo-
carpus ? sp., Lepidodendron cf. veltheimianum Sternb., and Bothrodendron australe
Feistm.

Rhacopteris semicircularis was first described by Lutz (1933 : 144) from the Culm
of Geigen near Hof. He compares it with R. circularis Walton, but considers it to
be different. Frenguelli's figures (pi. i, fig. i ; text-fig, i) show the specimen which
he compares with this species. He also compares it with Cardiopteridium, a com-
parison which is quite possible.

It is a pity that he did not figure his Rhacopteris ovata which he compares with
Steinmann's Rhacopteris circularis Walton.

The specimen which Frenguelli (1941, text-fig. 2) compares with Eremopteris
sanjuanina Kurtz resembles somewhat the specimens described later in this paper
as Triphyllopteris. A revision of the Argentine plants will possibly show more
comparisons with the Peruvian flora.

This is also the case with Eremopteris cf. whitei Berry (Frenguelli, 1941, text-fig. 3).
His determination may be right, and if so, this is the first time that Berry's species
has been recorded outside Peru.

The figure (pi. i, fig. 2) which Frenguelli compares with Lepidodendron veltheimi-
anum is indeterminable. There is no reason to accept a relation with Lepidodendron.
Here we have another example of the well-known fact that so many of the specimens
identified or compared with this, itself rather doubtful, species are absolutely value-
less. The same is true for the specimen figured in his pi. i, fig. 3, which he names
Bothrodendron australe Feistm. and that in pi. 2 which he calls Lepidodendron sp.

LIST OF THE CARBONIFEROUS FLORA OF PERU ACCORDING
TO THE OLDER LITERATURE

As a preliminary result of this review of the older literature on the Carboniferous
plants of Peru the following list of species is established :

Calamites cf. undulatus Sternb. (perhaps better as Calamites sp.). C. suckowi
Bgt. (Berry, 1922) ; C. peruvianus Gothan (1928) ; ? C. peruvianus Read
(1938) ; C. sp. Steinmann (1929, fig. 28).

202 THE CARBONIFEROUS FLORA OF PERU

Cyclostigma pacifica (Steinm.) Jongmans.

? Bothrodendron pacificum Steinmann (1929) ; Lepidodendron sp. Seward
(1922) ; Sigillaria or Lepidodendron sp. Seward (1922) ; Bothrodendron sp.
Seward (1922) ; " Bothrodendron " pacificum Gothan (1928).

" Lepidodendron " lissoni Steinmann (still doubtful).

Lepidodendropsis peruviana (Gothan) Jongmans.

Lepidodendron peruvianum Gothan (1928) ; Read (1938) ; Lepidodendron
obovatum Bgt. (Berry, 1922) ; Lepidodendron rimosum Sternb. (Berry,
1922, excluding fig. 3).

Asolanus (?) minimus Gothan (still doubtful).

Rhodea sp. Steinmann.

Rhodea filifera Steinmann (1911) ; Planta incertae sedis Seward (1922).

Sphenopteris paracasica Gothan.

Palmatopteris furcata (Bgt.) Berry (1922) ; Sphenopteris sp. Seward (1922) ;
Sphenopteris " parasica" Read (1938).

Sphenopteris whitei (Berry) Jongmans.

Eremopteris whitei Berry (1922) ; Sphenopteris hartlebeni Dkr. (Fuchs, 1900) ;
? Sphenopteris affinis (Steinmann, 1911) ; Adiantites whitei (Berry) Read
(1938).

Rhacopteris cf. circularis Walton.

Rhacopteris circularis Walt. (Gothan, 1928) ; Frenguelli (1943).

Rhacopteris ovata (McCoy) Walkom.

Rhacopteris circularis Walt. Steinmann (1929) at least very probable for the
crenulate margin ; Eremopteris peruianus Berry (1922, pi. 2, fig. 3 and
probably fig. 2) ; Rhacopteris ovata (McCoy) Frenguelli (1943).

Rhacopteris cf. R. cuneata Walkom.

Rhacopteris cf. cuneata Walk. (Read, 1938).

Rhacopteris peruiana (Berry) Jongmans.

Eremopteris peruiana Berry (1922, pi. 2, fig. I only) [not Adiantites peruianus
(Berry) Read (1938)].

THE CARBONIFEROUS FLORA OF PERU 203

cf. Triphyllopteris collombiana (Schimper).

Probably Adiantites peruianus (Read, 1938, text-fig. 3).
Possibly Adiantites bassleri Read (1938, text-fig. 7).

Aphlebia australia Read (1938).

Berry's Lepidostrobus (1922, pi. i, fig. 4) resembles a cone of Cyclostigma kiltor-
kense figured by Johnson (1913, pi. 41, figs. 3, 4), and Gothan's Trachyphyton negle-
gibile is very much like Johnson's figures of the rhizome of the same plant.

The age of the flora is considered to be Lower Carboniferous by Steinmann,
Seward (and Kidston), Gothan and Frenguelli. The only plant which could indicate
a Namurian or Westphalian age is Catamites cf . undulatus. However, the specimens
of Catamites recorded from Paracas are almost always indeterminable. Only in
one case does it seem to resemble in some respects Calamites undulatus.

Berry (1931 : 295) considers the flora as of Westphalian age, and if the floral
list given by him could be accepted, he certainly would be right. However, there
are many mistakes and wrong identifications in his list, and the revised list of the
specimens described by him definitely indicates a Lower Carboniferous flora.

The arguments of Gothan, Seward (Kidston), Steinmann, Read, and Frenguelli
are so weighty that there can be little doubt as to the lower Carboniferous age
of the flora. The description of the new collections will supply further evidence in
favour of their arguments.

A NEW COLLECTION FROM PARACAS

Many years ago the Trustees of the British Museum, by the kind intermediance
of Mr. W. N. Edwards, placed a new collection from this Peninsula at my disposal
for study. The plants were collected by the late Professor J. W. Gregory, whose
untimely death by drowning in a canoe accident in the rapids of the river Urubamba
at the age of 68 occurred, during an expedition to Peru, on 2nd June, 1932.

Sphenopteris whitei (Berry) Jongmans
(PI. 17, figs. 1-4)

? 1900. Sphenopteris hartlebeni Dunk. : Fuchs, p. 50.
? 1911. Sphenopteris affinis L. & H. : Steinmann, p. 50.

1922. Eremopteris whitei Berry, p. 20, pi. 4.

1938. Adiantites whitei (Berry) Read, p. 401.

? 1940. cf. Rhodea hochstetteri Stur : Jongmans & Koopmans, p. 228, pi. 4, fig. 9.
? 1941. Eremopteris cf. whitei Berry : Frenguelli, p. 470, text-fig. 3.

OCCURRENCE. Paracas, Peru ; Agua de los Jejenes, Argentina ; Rhas Gharib,
Egypt.

The collection includes a rather large number of specimens containing fragments
of a plant which may be determined as Sphenopteris whitei (Berry). Some of them
are figured here to prove their identity with those figured by Berry (1922) as Ere-

2o4 THE CARBONIFEROUS FLORA OF PERU

mopteris. There is no doubt about the agreement. The specimens do not show
any remarkable or new details. The form of the pinnules agrees with Berry's
figure. The venation, however, is not very clear. Amongst the numerous figures
with which these fragments may be compared is Rhodea hochstetteri Stur (Jongmans
& Koopmans, 1940, pi. 4, fig. 9) from the Lower Carboniferous of Egypt which is
almost certainly identical with Berry's species. Better examples of the Egyptian
material (collected more recently) will be described in a separate paper.

Other specimens belonging to this species are : ¥.25913, ¥.25934, ¥.25935,

V. 25937-39, ¥.25941-47.

<

Cyclostigma pacifica (Steinmann) Jongmans
(PI. 17, figs. 5-7 ; PI. 18, figs. 8-10 ; PL 19, figs. 11-14 i PI- 20> n§s- I4^2»3)

1922. Lepidodendron sp. Seward, p. 280, pi. 13, figs. 4-6.

1922. Sigillaria or Lepidodendron sp. Seward, p. 280, pi. 13, figs. 7, 8.

1922. Bothrodendron (?) sp. Seward, p. 281, pi. 13, fig. 9 ; text-fig.

1928. Bothrodendron pacificum Steinmann : Gothan, p. 296, pi. 13, figs. 3, $a.
cf. 1928. Lepidodendron sp. Gothan, p. 295, pi. 15, fig. 3.

1929. Bothrodendron pacificum Steinmann, p. 31, text-fig. 27.

Small branches with very approximate prominent leaf-cushions with a some-
what elliptical leaf-scar without cicatricules. Ligule not visible. Distance between
the cushions very variable, even on the same specimen. Cushions separated by
vertical, sharp, undulated lines, so that there is always an open communication
between the succeeding cushion-fields in the vertical row. Scars rounded in the
upper half, lower half consisting of two lateral sides meeting in a sharp point. In
older stems the distance between the scars is much greater, and the contours of the
cushions disappear as the distance increases. Surface of older stems smooth or
ornamented with undulating lines. Leaves long, narrow, and sharply pointed,
with a distinct midrib (Seward, 1922, pi. 13, figs. 4, 6).

DESCRIPTION. The collection contains numerous branches and small and large
stems. Several of the smaller stems belong to the same type and can be compared
with those figured by Seward (1922, pi. 13, figs. 4, 7, 8) and by Steinmann (1929,
text-fig. 27).

Unfortunately most of the specimens do not show the true leaf-scars as the pro-
minent cushions are always broken at the tips. There are, however, strong indi-
cations that these stems bore somewhat elliptical scars. The lines separating
the cushions are undulate but they are not always well marked, and in some cases
the form of the cushions is irregular and crushed (see Seward, 1922, pi. 13, figs. 7, 8).
This can be seen in PL 17, figs. 5, 6. PL 17, fig. 5«, is interesting as it shows the form
of the leaf-scar very clearly, and this is also visible on specimen ¥.25929. PL 17,
figs. 6, 7 and PL 18, figs. 8, 8a, show that the distance between the leaf-cushions is
very variable, and this is even more so in Steinmann's text-fig. 27 which also shows
that there are areas on the branches where the distance is much smaller than usual.
Specimens ¥.25922, ¥.25924, ¥.25926 and ¥.25927 all belong to this type but
are not so well preserved.

THE CARBONIFEROUS FLORA OF PERU 205

At first sight it seems somewhat peculiar that the leaf-cushions figured by Seward
(1922, pi. 13, figs. 7, 8) belong to the same plant as his fig. 4. However, PI. 17,
fig. 5, and PI. 18, fig.8, show that both types are present on the same specimen.
It is possible that there are irregularities in the form of the cushions such as those
seen in Sigillaria in regions where the large scars bearing the fructifications are
found, or it may be that there is some difference between succeeding regions of stems
and branches as is known from other species of Sigillaria in connexion with differences
in the mode of growth (cf. Potonie, 1894). A similar difference is also present in
Bothrodendron leslii Seward, in Sigillaria mutans Weiss, and in Steinmann's text-
fig. 27.

One of the best specimens, with part of its counterpart, is shown in PI. 18, figs. 9,
10. It shows stems on both sides. The branch in fig. 9 can be compared with
Seward's fig. 4. In the photograph the cushions are not always distinctly visible,
but the undulated lines between the vertical rows are well seen in several
places.

The true scars are well preserved on the counterpart (PL 18, figs. 10, ioa). The
separation of the rows of cushions is indicated by a very delicate undulated line,
so that the leaf-cushions are situated in the centre of the broadest part which is
narrowed up- and downward. There is always an open communication between
the succeeding cushion-fields in the vertical rows. The real scar lies in a deep
hollow of the impression (in reality an elevation). Over and under the scars and
cushions there is a distinct smooth field. There is no special ornamentation on the
rest of the cushions. These characters are very well seen in PL 17, fig. 5. No
cicatricules or ligule are visible.

As already stated the space between the cushions and scars is variable. In PL 17,
fig. 7, the space is considerable. At the same time the lines separating the vertical
rows are less distinct and disappear, the characters of the cushions also decrease,
and the more or less isolated scars are all that remains of the former structure. The
form of the scar itself does not change very much ; it is somewhat more regularly
elliptical. This is well seen in PL 19, figs, n, 12. and in the counterpart specimen
(V. 25933). A very good example of this type is shown in PL 19, fig. 13 ; here the
form of the leaf-scars is clearly visible. They have about the same form as those
seen in PL 17, fig. 5 and PL 18, fig. 10. Another good example is given in PL 19, fig.
14, which can be compared with Seward's pi. 13, fig. 9.

That the isolated scars originally had the same form as that shown in PL 18,
fig. 8 can be seen on the right of PL 19, fig. n, where the elongated cushions and the
undulating lines which separate them are visible.

The surface between the scars is smooth (or almost smooth) in the specimen
figured by Seward (1922, pi. 13, fig. 9) and in that figured in PL 19, fig. n. However,
the specimens shown in PL 19, figs. 12, 14 are remarkable for a distinct ornamenta-
tion on the surface between the scars. This ornamentation consists of delicate
longitudinal lines, a number of which converge towards the scars, and recalls that
of Bothrodendron. As the ornamentation is not always visible it may be that its
presence or absence is due to the layer which is preserved on the fossil. The
ornamentation is somewhat similar to that figured by Gothan (1928, pi. 15, figs, za,

GEOL. II, 5. I7

206 THE CARBONIFEROUS FLORA OF PERU

3) in his Asolanus (?) minimus. In the British Museum specimens the arrangement
is visible in oblique lines and less distinctly in horizontal lines.

It is clear that, as in the case of the Westphalian species of Bothrodendron,
Cyclostigma kiltorkense, and some species of Lepidodendropsis, the leaf-cushions on
the young branches are very distinct, approximated, and more or less separated by
dividing lines, so that they have a lepidodendroid aspect. On the lower parts
of the branches and on the stems this lepidodendroid character decreases and dis-
appears. The result is that the leaf-scars are placed on a rather smooth or delicately
ornamented surface (Seward, 1922, pi. 13, fig. 9). This stage is seen in the specimen
(with its counterpart) shown in PL 19, figs. 12, 14. An excellent specimen of this
type is figured in PL 19, fig. 146. One part of the specimen shows a not very distinct
ornamentation between the scars, another part is almost smooth but shows the scars
very well. The scars have the same form as those in PL 17, figs. 6, 7, but are much
better preserved. The specimen represented in PL 19, fig. n, is, in some respects,
transitional between the extremes.

Rarely the leaves are still attached to the branches. The best examples are
shown in Seward's pi. 13, figs. 4, 6 ; the leaves are long, narrow and sharply pointed,
with a distinct midrib. The specimens show fragments of leaves rarely or not at
all.

This species must belong to Cyclostigma and not to Bothrodendron, for the form
of the leaf-cushions and their arrangement on the younger branches are entirely
different from the latter and no ligule or cicatricules can be found. Steinmann's
specific name pacificum must be retained, and therefore the correct name for these
plants is Cyclostigma pacifica (Steinmann) Jongmans.

Specimens similar to those figured in PL 19, figs. 12, 14, with ornamentation on
the surface between the leaf-scars, may be distinguished as var. decorata, although
as stated above the presence or absence of this ornamentation may be due to
preservation.

Regarding the determination of this plant as Cyclostigma, the " 'Lepidostrobus "
figured by Berry (1922, pi. I, fig. 4) and the peculiar specimen described and figured
as Trachyphyton neglegibile by Gothan (1928, pi. 14, figs. 3, 4) may be important.
The Lepidostrobus can be compared with the figures of cones of Cyclostigma kiltorkense
in Johnson's paper (1913, pi. 14, figs. 3, 4) and Gothan's figures with those of the
rhizome of this plant figured by the same author (Johnson, 1914, pis. 14, 15). It
may yet be proved that these or similar organs belong to Cyclostigma pacifica or
an allied species.

Genus LEPIDODENDROPSIS Lutz

Lepidodendropsis peruviana (Gothan) Jongmans

(PL 20, fig. 15)

1922. Lepidodendron rimosum Sternb. : Berry, p. 24, pi. 8, figs, i, 2 only.

1922. Lepidodendron obovatum Brongn. : Berry, p. 26, pi. i, fig. 5.

1928. Lepidodendron peruvianum Gothan, p. 294, pi. 13, fig. 2.

1929. Lepidodendron peruvianum Gothan : Steinmann, p. 30, text-fig. 23A-D.
1938. ? Lepidodendron peruvianum Gothan : Read, pp. 398, 402, text-fig. 4.

THE CARBONIFEROUS FLORA OF PERU 207

DESCRIPTION. As stated above the specimens identified as Lepidodendron peru-
vianum by Gothan (1928) and by Steinmann (1929) belong to Lepidodendropsis.
Three examples of this species are in the British Museum collection (¥.25911,
V. 25912 and V. 25931). The best specimen is that seen in PI. 20, fig. 15. Form
and arrangement of the cushions agree with those figured by Gothan (1928, pi. 13,
fig. 2) and by Steinmann (1929, text-fig. 23). There is, however, a slight difference.
According to their figures and descriptions the surface of the bands separating the
leaf-cushions is smooth. This is not the case in the new specimens, where some
rather coarse longitudinal or almost longitudinal lines are present. This difference
may be essential or may be due to the state of preservation. At all events, the
comparison made between this species and the Egyptian Lepidodendropsis fenestrata
Jongm. & Koopm. increases in probability.

Lepidodendropsis ("Lepidodendron") lissoni (Steinmann) Jongmans

(PL 20, fig. 16)

1928. Lepidodendron lissoni Steinmann : Gothan, p. 295, pi. 14, fig. 2.

1929. Lepidodendron lissoni Steinmann, p. 31, text-fig. 24.

DESCRIPTION. The British Museum collection contains a specimen (¥.25932)
which may belong to the same species as that figured by Gothan (1928) and by
Steinmann (1929). Gothan compares the species with Lepidodendron spetsbergense
Nath. The specimen is distinguished by the peculiar field under the scars. It
may be that such fields are the remains of lepidodendroid cushions, and it is probable
that this plant belongs to Lepidodendropsis, in which case it might be compared
with Lepidodendropsis cyclostigmatoides Jongmans, Gothan & Darrah (1937, pi. 50,
fig. 28). Since no true Lepidodendron has been found in the flora of Paracas it is
safer to use the name Lepidodendropsis ? lissoni (Steinmann) Jongmans. Better
specimens are necessary before a final decision is possible.

? Lepidodendropsis sp.

(PI. 20, fig. 17)

This remarkable specimen (¥.25915) shows two large branches or stems, at least
7-8 cm. broad. They are covered with cushions very similar to those of Lepidoden-
dropsis. No real leaf-scar is visible. One of the stems shows fragments of the leaves
attached to it, and it is clear that in this species the leaves were very persistent.
Although it is almost certain that the specimen belongs to an undescribed species,
the details are too poorly preserved for specific determination.

Rhacopteris cf. cuneata (Walkom)

(PI. 20, fig. 18)
1938. Rhacopteris cf. cuneata (Walkom) : Read, p. 401, text-fig. 5.

Two small specimens (counterparts, ¥.25948) can be compared with Rhacopteris
sp. cf. R. cuneata (Walkom) figured by Read (1938, text-fig. 5). The specimens are

2o8 THE CARBONIFEROUS FLORA OF PERU

illustrated in PI. 20, figs. 18, i8a, i8av The occurrence of a very small fragment of
Rhacopteris ovata, which can be seen at the top left corner of PL 20, fig. i8#, is
interesting. The only previously recorded specimen of Rhacopteris ovata from
Paracas is that figured by Berry (1922, pi. 2, figs. 2, 3) as Eremopteris peruviana.
Better examples of this species are known from Vichaicoto, S. of Huanuco, Peru
(Steinmann, 1929, text-fig. 29) and from Peru and Argentina (Frenguelli, 1943,
pis. 1-4).

It is curious that the present collection does not contain specimens of Rhacopteris
circularis nor, except for the fragment referred to, of Rhacopteris ovata which must,
however, belong to the more common elements of the Peruvian flora.

Asolanus (?) minimus Gothan
1928. Asolanus (?) minimus Gothan, p. 295, pi. 15, figs. 2, 2#, ? 3 right.

The collection contains two small fragments (V. 25909 and ¥.25910) which may
be compared with Gothan's species. Unfortunately the preservation is too bad
for the specimens to be figured.

From the flora represented in the British Museum collection it follows that although
the number of species is relatively small, it provides more and better evidence
regarding several of these species.

The full list of the flora so far known from the Paracas Peninsula contains the
following species :

Catamites sp. (cf. undulatus Sternb.).

Cyclostigma pacifica (Steinm.).

Lepidodendropsis peruviana (Gothan).

Lepidodendropsis sp.

Lepidodendropsis ? lissoni (Steinm.).

Asolanus ? minimus Gothan.

Rhodea sp.

Sphenopteris paracasica Gothan.

Sphenopteris whitei (Berry).

Rhacopteris ovata (McCoy).

Rhacopteris ? circularis Walton.

cf. Rhacopteris cuneata (Walkom).

cf. Triphyllopteris collombiana (Sch.)

Adiantites bassleri Read (Triphyllopteris).

Aphlebia australis Read.

" Lepidostrobus " sp. Berry (? Cyclostigma).

Trachyphyton neglegibile Gothan (? Stigmaria, ? Cyclostigma).

The British Museum collection provides further proof that the flora of Paracas
must be of Lower Carboniferous age. The aspect of the flora is similar to that of
the Lower Carboniferous of Egypt and of the Pocono in U.S.A., although there are

THE CARBONIFEROUS FLORA OF PERU 209

minor differences ; thus Rhacopteris of the Peruvian type is not yet known from
Egypt or the Pocono formation.

FLORA OF CARHUAMAYO

Dr. De Voogd, one of my former assistants, sent me a good collection of plants
from the neighbourhood of Cerro de Pasco at Carhuamayo. With the exception
of the specimens of Rhacopteris figured by Frenguelli (1943) no descriptions or
figures of plants from this locality have so far been published. Read (1941 : 17)
mentions a florule from this locality containing Rhacopteris ovata (McCoy), Adiantites
bassleri Read (possibly identical with his figure (1938 : 402, text-fig. 7) from Paracas,
belonging to Triphyllopteris), and Lepidodendron pernvianum Gothan (erroneously
ascribed by the author to Berry & Read) which probably belongs to Lepidodendropsis
peruviana (Gothan).

The new collection is important for the remarkable forms of Lepidodendropsis
and the very good specimens of Rhacopteris and Triphyllopteris.

Lepidodendropsis de voogdi n. sp.

(PI. 21 ; PI. 22, figs. 24-27 ; PI. 25, fig. 32 ?)

DIAGNOSIS. Stems with leaves, sometimes with sporangia. Stems with scars
of Lepidodendropsis type, covered by the basal parts of the leaves. Scars numerous.
Distance between scars small. Leaves spreading, 3 cm. or more long, up to 5 mm.
broad, acuminate, with a distinct middle nerve. Sporangia, containing megaspores,
visible in the axils of the leaves.

DESCRIPTION. PI. 21, fig. 19 shows the axis and leaves (or sporophylls) of a
specimen which make a somewhat lepidodendraceous impression. The axis is very
broad. The scars, badly preserved, are placed in obliquely ascending lines. The
horizontal and vertical arrangements are both distinct, but the oblique arrangement
is most striking. Numerous leaves (or sporophylls) are connected with the axis.
They are 3 cm. or more long and up to 5 mm. broad, acuminate, with a distinct
middle nerve. The lower (sporangium-bearing) part is rather short, about 5 mm.
long, and increases in breadth towards the implantation of the axis. This basal
part is not very clearly separated from the rest. The leaves are spreading and very
crowded. They resemble a Lepidophyllum of the lanceolatum type.

A second specimen (PI. 21, fig. 20) is much longer. It shows the sporophylls,
but the form of these is not so well seen. The most interesting part of this specimen
is where the large, Lepidostrobus-\ike sporangia are visible on the outer left and right
sides. They are 3-4 mm. broad and about 2 mm. high. Their size agrees with
the basal part of the sporophylls in the first specimen. The specimen shows that
the sporangium-bearing part of the sporophyll is almost horizontal and that the free
part is abruptly erect.

The form of the scars on the broad axis is not visible, probably due to fragments
of the sporophylls which more or less cover them. As far as can be seen, they
appear to be almost elliptical.

aio THE CARBONIFEROUS FLORA OF PERU

A specific determination of the strobilus, or perhaps sporophyll bearing branches
is of considerable importance. In this respect a specimen with similar leaves, but
with no sporangia present (PL 21, figs. 21, 210) is of value. It shows two leaf-
bearing branches, one of which, the branch on the left (PL 21, fig. 2ia), shows scars
which agree completely with the small scars of Lepidodendropsis .

Another branch shows a bifurcation (PL 21, figs. 22, 220). The scars are oval
elliptical and are about the same size as those shown in PL 21, figs. 21, 2ia. The
leaves are broken. In the axils of the leaves are the remains of sporangia which
contain megaspores.

This specimen is clearly a fructification which does not form a true strobilus and
which agrees in this respect with Lepidodendropsis vandergrachtii. The latter
species differs, however, as in Pinakodendron, in the absence of lepidodendroid
sporangia which are present in the Peruvian material. On the back of this specimen
two fragments of branches are visible both with long leaves attached (PL 21, fig. 23).
One, on the left, without, the other, on the right, probably with sporangia and spores.

Another specimen (PL 22, fig. 24) shows two fragments of branches, both with well-
preserved leaves and probably sporangia and spores. The leaves show the general
form of those in PL 21, fig. 19, but they appear to be somewhat narrower. PL 22,
fig. 25, shows a narrow branch with very long leaves of the same type. There is no
trace of sporangia or spores.

A somewhat curious specimen (PL 22, fig. 27) shows leaves only without a trace
of the branches or stems. The specimen figured in PL 22, fig. 26 shows isolated
leaves which are unusually narrow.

This plant undoubtedly belongs to Lepidodendropsis. It is here named Lepidoden-
dropsis de voogdi after Dr. De Voogd who collected the material for me.

Lepidodendropsis cf. de voogdi Jongmans
(PL 25, fig. 32)

This specimen has no leaf-scar and the " cushion " is of the Lepidodendropsis
type. It is possible that it belongs to one of the other species, but it may be different.
It can be compared with Lepidodendropsis de voogdi (PL 21, fig. 21) which shows the
same type of leaf-bases.

Lepidodendropsis steinmanni n. sp.

(PL 22, figs. 28a, d; PL 23, figs. 286, c ; ? PL 24, fig. 31)

DIAGNOSIS. Large stem, more than 3-5 cm. broad, covered by small " cushions."
Horizontal distance between two cushions about 0-8 cm., oblique distance 4-5 mm.
Vertical distance variable. Cushions occasionally more crowded, very small and
not very distinct, indicated by smooth, almost triangular spaces (probably represent-
ing the basal parts of the leaves attached at the top of the " cushions "). Leaves
attached at the " cushions " with their full base ; on leaving the stem they are first
directed downwards and then obliquely erect.

THE CARBONIFEROUS FLORA OF PERU 211

DESCRIPTION. A second species of Lepidodendropsis is represented by a large
stem with counterpart. Between these two parts a fragment of the pith cast is
present (PL 22, fig. z8d). The stem is rather broad, somewhat more than 3-5 cm.
wide and covered by small " cushions " (PL 22, fig. 2,80) arranged in ascending lines.
The arrangement in horizontal lines is not very distinct. The horizontal distance
between two cushions is about 0-8 cm., the oblique distance 4-5 mm. The distance
between the scars is not always the same. In at least two places they are more
crowded, as in Lepidodendropsis vandergrachtii from the Pocono. The best
"cushions " can be seen at the top of the figure. They are very small and agree
generally with those of Lepidodendropsis (PL 23, fig. 28c) ; a cushion is indicated
by a smooth, almost triangular space. A leaf-trace is not visible, and the upper
part of the " cushion " bends into the rock over the upper limit of the " cushion."
The space between the cushions is finely granulated (chagrinate) and almost smooth
(PL 23, fig. 28c).

The pith-cast is covered by a thin coal-layer which is somewhat more distinctly
chagrinate than the impressions. This delicate ornamentation is also visible on
the actual surface of the pith-cast (PL 22, fig. 28d).

The stem bears narrow leaves which are specially well seen on the upper part.
The leaves are directed downwards for about 4 mm. as they leave the stem, but
from there they are abruptly erect.

This species agrees rather well with Cyclostigma ungeri Jongmans, Gothan &
Darrah (1937, pi 57, figs. 45, 45#). Here too the horizontal lines in the arrangement
of the cushions are much less developed than the very distinct oblique lines, and the
surface bears a very delicate ornamentation. This ornamentation is different,
however, and the cushions are larger, not so punctiform, and the horizontal and
vertical distances between the cushions are much greater.

The leaf-cushions of the Peruvian specimens do not agree with those of Cyclostigma
and more closely resemble those of Lepidodendropsis. A true leaf -scar is not visible.

In the description of Cyclostigma ungeri it was pointed out that it was only
provisionally compared with that genus. It probably belongs to Lepidodendropsis.

The new name Lepidodendropsis steinmanni is provisionally proposed for the
Peruvian species.

Lepidodendropsis cf. steinmanni Jongmans
(PL 24, fig. 31)

In this specimen the leaf-cushions are similar in form to those of Lepidodendropsis
steinmanni but they are much larger. It may be an older stem of this species and
is named Lepidodendropsis cf. steinmanni accordingly.

Cyclostigma pacifica (Steinmann)
(PL 24, fig. 30)

A small specimen belonging to this species may be compared with those figured by
Seward (1922, pi. 13, figs. 4, 6). It shows the shape of the cushions and also

212 THE CARBONIFEROUS FLORA OF PERU

fragments of the leaves. It is remarkable that this is the only specimen in the
new collection which undoubtedly belongs to this species.

Cyclostigma cf. pacifica (Steinmann) var.
(PI. 23, fig. 290 ; PL 24, fig. 296, c)

A plant which probably belongs to Cyclostigma is represented by a small fragment
found in the same block as the specimen figured in PL 22, fig. 28. It consists of the
impression of the upper and lower sides of the stem and a part of the pith-cast
(PL 23, fig. 29^ ; PL 24, fig. 296, are from the upper and lower surfaces, PL 24,
fig. 29C, from the pith-cast).

The leaf-cushions have the same form as those found in Cyclostigma pacifica.
The only difference is that they are much more flattened, and are not so prominent
as in most of the specimens of that species. The surface of the cushions is not
smooth. There are also fragments of leaves which are shown in PL 24, fig. 296, c.

A detailed description seems unnecessary since the figures show the details rather
well. The lines separating the cushions are delicate but sharp and are well seen in
several places (PL 23, fig. 290, left side; PL 24, fig. 296). The leaf-scars are placed
in the centre of the cushions. The surface is finely chagrinate.

It is very probable that this plant is specifically identical with Cyclostigma pacifica
and it is therefore described as C. cf. pacifica var. until more complete specimens
are available for study.

Rhacopteris ovata (McCoy) Walkom
(PL 25, figs. 33-37 ; PL 26, fig. 45)

1922. Eremopteris peruianus Berry, p. 19, pi. 2, fig. 3, ? fig. 2.
1929. ? Rhacopteris circularis Walton : Steinmann, p. 33, text-fig. 29 A-C.
1938. Rhacopteris ovata (McCoy) : Read, p. 401, text-fig, i.

1943. Rhacopteris ovata (McCoy) : Frenguelli, pp. 14, 22, pi. i ; pi. 2, fig. i ; pi. 4, figs. 1-3
(Argentina) ; PI. 3, fig. 2 (Carhuamayo) .

OCCURRENCE. Paracas ; Vichaicoto, S. Huanuco and Carhuamayo, Peru. El
Tupe, La Rioja, Argentina.

As in Vichaicoto, S. of Huanuco, specimens of Rhacopteris of different sizes are
very common at this locality. Some of them are excellently preserved.

In many cases it is not easy to distinguish the two species Rhacopteris circularis
and Rhacopteris ovata. The chief differences are the margins and the form of the
pinnules. In typical specimens the margins of Rhacopteris ovata are crenulate
and the form of the pinnules is distinctly asymmetric and not circular. However,
in Walton's description of Rhacopteris circularis (1926 : 208) he states " Margin of
pinnules entire, more rarely crenulate or lobed. Pinnules circular to semi-flabelli-
form. There is a tendency to asymmetry in the higher pinnules on the frond."
It is possible that in most cases the smaller specimens show the characters of R.
circularis and the larger ones those of R. ovata. Frenguelli (1943) figures specimens

THE CARBONIFEROUS FLORA OF PERU 213

which he ascribes to both species. His figures of R. ovata (pi. i ; pi. 3, fig. 2 ; pi. 4,
figs. 1-3, especially fig. 2) show the characters of R. ovata very well, with crenulate,
asymmetric large pinnules.

Frenguelli's pi. 3, fig. i, shows rounded pinnules and no crenulation, whereas
the specimen in pi. 4, fig. 4, shows pinnules even more rounded and slightly asym-
metric, also without crenulation. In some respects this latter specimen is transi-
tional between the two extreme types.

Berry's figures (1922, pi. 2, fig. 3 and possibly fig. 2) show the characters of R.
ovata. Steinmann's figures (1929, text-figs. 2ga-c) can also be identified with this
species, but Gothan's figure (1928, pi. 15, fig. i) is somewhat doubtful ; most of
the pinnules are incomplete and it is possible that they may have been more rounded.
As far as can be seen there is no crenulation. The specimen figured by Read (1938,
text-fig, i) also shows the characters of R. ovata.

Amongst the figures of R. ovata in the older literature is an interesting specimen
figured by Dun (1905, pi. 23). This specimen is very long, the lower part of the
pinna shows the large, crenulate and more or less asymmetric pinnules typical of
R. ovata, whereas the upper part of the pinna shows much smaller symmetric
pinnules with distinctly circular upper margins. Similar small pinnules are also
represented in Dun's pi. 22, fig. 3.

Several of the specimens figured by Feistmantel (1890) do not show the crenulated
margin and some of them are more or less circular (e.g., specimens in pi. 4), whereas
others agree more closely with R. ovata (pi. 5, fig. 2 ; pi. 8 ; pi. 9, especially fig. 2).
Some crenulation is indicated in his pi. 7, fig. i.

Walton's figures of R. circularis generally show a uniform character in the shape
of the pinnules, which is never or very rarely found in specimens of R. ovata. There-
fore I am inclined to consider most of the Peruvian specimens as belonging to R.
ovata.

In most of the specimens the margins of the pinnules are not preserved or have been
broken off or hidden in the rock. This is the case with much of the present material,
although, on most of these specimens, the crenulation is visible somewhere. The
shape of the pinnules differs considerably. An extreme type is the large specimen
shown in PI. 25, fig. 33. A number of specimens (PI. 25, figs. 34-37) show pinnules
of different sizes, and some of them show the venation quite well.

Rhacopteris cf. circularis Walton
(PL 25, fig. 38 ; PL 26, fig. 39)

1928. Rhacopteris circularis Walton : Gothan, p. 293, pi. 15, fig. i.

1943. Rhacopteris circularis Walton : Frenguelli, pp. 22, 41, pi. 3, fig. i ; pi. 4, fig. 4.

OCCURRENCE. Carhuamayo and Vichaicoto, Peru. Agua Salada, La Rioja,
Argentina.

A fine specimen of Rhacopteris is represented in PL 25, fig. 38. Here most of the
pinnules have more or less rounded margins and their dimensions are very variable.
It is interesting to note that the pinnules show a distinct footstalk, as in Walton's

2i4 THE CARBONIFEROUS FLORA OF PERU

figures of RhacopUris circularis. If R. circularis is present in the Peruvian flora,
then this specimen may be considered to belong to it. Feistmantel's figured speci-
mens with margins and pinnules typical of Rhacopteris ovata do not possess such a
distinct leaf stalk. It is possible that the specimen figured in PI. 26, fig. 39, which
has rather distinct leaf stalks also belongs to Rhacopteris circularis Walton.

Genus TRIPHYLLOPTERIS Schimper

This genus is well represented in the collection from Carhuamayo. Three forms
can be distinguished and they may belong to different species.

Triphyllopteris collombiana (Schimper)
(PL 26, figs. 40-42)

? 1938. Adiantites peruianus (Berry) Read, p. 401, text-fig. 3.
? 1938. Adiantites bassleri Read, p. 399, text-fig. 7.

1941. cf. Eremopteris cf. sanjuanina Kurtz : Frenguelli, p. 468, text-fig. 2.

OCCURRENCE. Paracas and Carhuamayo, Peru ; Agua de los Jejenes, Argentina.

A rather large form is represented by several specimens of which PL 26, figs. 40-42,
are typical examples. These specimens resemble Triphyllopteris collombiana as
figured by Schimper (1862, pi. 25, figs. 8-10, Cyclopteris] from Burbach. The
nervation is well shown in PL 26, fig. 42.

It is possible that the specimens figured as Adiantites peruianus and A. bassleri
by Read (1938) belong to this species.

Triphyllopteris lescuriana (Meek)
(PL 26, figs. 43-45)

The specimens referred to this species are much smaller and the leaves are more
divided than in Triphyllopteris collombiana. In every case the pinnules are divided
in a way which is typical for this genus.

The specimens represented in PL 26, figs. 43, 44, have elongated, acuminate
divisions of the individual segments. Fig. 43 is part of the top of a leaf and shows
the division of the segments very clearly. As far as can be seen the venation agrees
with that described for Triphyllopteris.

It is very probable that the specimen figured in PL 26, fig. 45 also belongs to this
species, although the divisions of the pinnules are not so sharply acuminated. Good
examples of Rhacopteris ovata occur on this specimen, one with large, and one with
small pinnules.

These specimens agree rather well with Triphyllopteris lescuriana (Meek) as
figured by Jongmans, Gothan & Darrah (1937, pi. 44, figs. 7, 8) and by Lesquereux
(1880, pi. 50, fig. 6).

THE CARBONIFEROUS FLORA OF PERU 215

? Triphyllopteris peruviana n. sp.
(PI. 26, fig. 46)

DIAGNOSIS. Dimensions much smaller than in T. lescuriana and the division
of the leaf much denser. Pinnules divided in five lobes, the top lobe elongate, the
side lobes rounded and much shorter than in T. lescuriana.

DESCRIPTION. The third form is represented by one good specimen (PI. 26,
fig. 46) and one fragment. The dimensions are much smaller and the division of
the leaf is much denser. The division of the pinnules into three lobes is not so regular.
In most cases there are five lobes, the top lobe being elongate, the side lobes more
rounded and much shorter. It is not certain that this specimen really belongs
to Triphyllopteris and it is therefore provisionally named ? Triphyllopteris peruviana.
More and better material is necessary before it can be attributed to this genus with
certainty.

CONCLUSION

The flora of Carhuamayo contains :

Lepidodendropsis de voogdi Jongmans.
Lepidodendropsis cf. de voogdi Jongmans.
Lepidodendropsis steinmanni Jongmans.
Lepidodendropsis cf. steinmanni Jongmans.
Lepidodendropsis sp.

Cyclostigma pacifica (Steinmann) Jongmans.
Cyclostigma cf. pacifica (Steinmann).
Rhacopteris ovata (McCoy).
Rhacopteris cf. circularis Walton.
Triphyllopteris collombiana (Schimper).
Triphyllopteris lescuriana (Meek).
? Triphyllopteris peruviana Jongmans.

It is clear that this flora, as was to be expected, is of Mississippian age.

THE DISTRIBUTION OF LEPIDODENDROPSIS

It is clear that the age of the Peruvian Carboniferous floras is Mississippian, and
this was the opinion of most of the earlier authors. Berry, however, accepted a
Pennyslvanian age for Paracas, but it has since been proved, in particular by Gothan,
that the determinations upon which Berry based his conclusion were not correct.
The records described in this paper fully agree with a Mississippian age.

As previously stated the Peruvian flora is very similar to that of the Pocono in the
United States. A number of species of Lepidodendropsis were described from the
Pocono (Jongmans, Gothan & Darrah, 1937) some of which resemble the type
species Lepidodendropsis hirmeri Lutz. Other species such as L. vandergrachti have
quite a different habit and are more like Sigillaria. The fructification of this species
is not a strobilus but, as in Pinakodendron, the spores are found at the base of small

216 THE CARBONIFEROUS FLORA OF PERU

sporophylls on young twigs. A similar fructification is recorded for the Peruvian
Lepidodendropsis de voogdi Jongmans. Other important elements of the Pocono
flora are some species of Triphyttopteris. Rhacopteris was not present in the Pocono
collections examined.

Trochophyllum breviinternodium (Arnold, 1933 ; renamed Prolepidodendron,
Arnold, 1939, pi. I, fig. 2) very much resembles Lepidodendropsis. The only differ-
ence is in the form of the leaves, with their broad, flattened, upper ends. It is
possible that this species is in some way a transition with Protolepidodendron Krejci
which is certainly related to Lepidodendropsis but distinguished by the leaves
which in their upper parts are divided into two. If the leaves are not preserved,
or the division of the leaves is not visible, it is almost impossible to distinguish the
stems of Protolepidodendron scharyanum, as figured by several authors, from those
of Lepidodendropsis.

Arnold's specimens are from the Upper Devonian (Oswayo sandstone), near
Port Allegany, McKean County, Pennsylvania, which was originally placed in the
Pocono (Arnold, 1933).

Prolepidodendron breviinternodium (Arnold) has been found associated with
Archaeopteris cf. roemeriana, Rhacopteris sp. (cf. R. circularis Walton from the Teilia
beds), and fragments of Callixylon.

Colpodexylon Banks (1944) is another related genus with dichotomous or three-
forked leaves.

Lutz (1933) described Lepidodendropsis as a new genus from the Lower Carboni-
ferous of Geigen near Hof, Bavaria. In that flora L. hirmeri is represented by an
abundance of very good specimens. Other important elements in the flora are :
Sphenophyllum saxifragaefolioides Leyh., 5. geigense Lutz, Neurocardiopteris, Cardi-
opteris, Rhacopteris lindsaeformis Bunb., R. semicircularis Lutz, Archaeopteridium
dawsoni Stur, Sphenopteridium, Calathiops and different species of Rhodea.

It is interesting to note that Lacey (1952 : 375,376) mentions two specimens from
the Lower Carboniferous of Wales both of which may well belong to Lepidoden-
dropsis. One of the specimens is from Craig Quarry, nr. Denbigh. The other
(p. 375) comes from the Dyserth locality and was originally named Lepidophloios
cf. laricinus Steinberg.

Dubertret (1933 : 288) has recorded lepidodendroid remains associated with a
Tournaisian fauna in the Jebel Abd el Aziz, North-east Syria, and Mr. W. N.
Edwards informs me that there are some fragments probably referable to Lepido-
dendropsis in the British Museum (Natural History) collected by W. A. Macfadyen
from the Wadi Gharra, Jebel Abd el Aziz.

Another flora which may be compared with that of the Peruvian Lower Carboni-
ferous was described from Egypt (Jongmans & Koopmans, 1940). It was collected
from samples of cores from wells in Rhas Gharib. The flora contains Lepidoden-
dropsis fenestrata J. & K., Sphenopteris whitei (Berry)- — originally named Rhodea cf.
hochstetteri ; the new determination, not yet published, is based on better specimens
collected in 1946 — and Cyclostigma aegyptiaca J. & K.

In the same paper some specimens from Wadi Um Shebba, Sinai, were figured,
which may also be compared with, and probably belong to Lepidodendropsis (Lepi-

THE CARBONIFEROUS FLORA OF PERU 217

dodendropsis sinaica J. & K. including two specimens of Lepidodendron mosaicum
Salter figured by Seward, 1932, pi. 21, fig. 4, and pi. 22, fig. 9). In connection
with the Carboniferous of Egypt it is interesting that Porodendron sp. described by
Gothan (1933) from the Oasis di Cufra, may also belong to Lepidodendropsis.

An important contribution to the Lepidodendropsis flora of North Africa was
published by Boureau (1954). He records the occurrence of Rhacopteris ovata and
R. circularis in the Lower Carboniferous (? Dinantien) of Air, near Tafadeck,
Central Sahara. Although Lepidodendropsis itself was not found at this locality,
it is very probable that the horizon is that of the Lepidodendropsis flora, for
according to a young French geologist whom I met in Algeria, he had collected
Lepidodendropsis from the Lowest Carboniferous of the Sahara. I have not yet,
however, seen the specimen which he promised to send to me for examination.

A plant which certainly belongs to Lepidodendropsis is Sigillaria fezzanensis
Chiarugi (1948). This specimen is much like Lepidodendropsis vandergrachti J.,
G. & D. from the American Pocono and probably belongs to this species. It has
nothing to do with Sigillaria and certainly not with 5. brardi Bgt. with which it is
compared by Chiarugi. On the strength of this comparison Chiarugi states that the
locality belongs to an elevated part of the Carboniferous. In my opinion it belongs
to the Lower Carboniferous or Upper Devonian in agreement with the marine
fossils, which according to Borghi (1939) indicate a transition between the Mississip-
pian and the Upper Devonian.

Chiarugi (1948 : 81) compares his material with Fritel's Archaeosigillaria vanuxemi
Goepp. (1925, pi. 3, figs. 1-5). These specimens have leaf-cushions which are
very approximate. However, fig. 3, upper part, and fig. 5 show clearly that the
space between the individual leaf-cushions can become much larger. It is not
clear from the figures whether there is a true leaf -scar, as in Archaeosigillaria, or not.
A comparison of Archaeosigillaria and Lepidosigillaria with Lepidodendropsis can
only be made by an examination of the original specimens.

Another figure in Fritel's paper with which comparison is possible is his Lepido-
dendron cf. volkmannianum (pi. 3, fig. 6b). This specimen was found at the same
locality as his Archaeosigillaria.

Carpentier (1930) described a small flora from the Lower Carboniferous of Morocco.
His Lepidodendron ? aff. corrugatum (pi. i, fig. 3), Epi de Calamariee ? (pi. 2, fig. 3)
and Lepidodendron veltheimi (pi. 4, figs. 1-3) most probably belong to Lepidoden-
dropsis. It may even be that his Arctodendron (pi. 5, figs. 2, 2 bis) is an old stem of
Lepidodendropsis. It is very curious, however, that at this locality, some very
poorly preserved specimens were collected which simulate impressions of Sigillaria.

Lepidodendropsis hirmeri has also been collected by Melendez and myself (1950)
from the Lower Carboniferous of Valdeinfierno, Spain. Here it occurs at the same
locality, but not exactly in the same place, with a flora containing Asterocalamites
scrobiculatus Schl., Stigmaria stellata Goepp., Sphenophyllum saxifragaefolioides
Leyh, 5. geigense Lutz, Rhodea cf. stachei Stur, R. cf. moravica Ett., Triphyllopteris
cf. minor J. & G., T. collombiana (Sch.), and Calathiops cf. plauensis Gothan.

This flora very much resembles that of Geigen, near Hof , and both may be compared
with the floras of the Pocono, of Peru and of Egypt.

2i8 THE CARBONIFEROUS FLORA OF PERU

In the Donetz Basin the Lepidodendropsis flora is said to occur in the Upper
Devonian. Zalessky (1931) described Heleniella theodori and compared it with some
species of Sigillaria (S. youngiana Kidston, 5. tschirkovaeana Zal., and 5. canobiana
Kidston) which show undulated ribs. However, an examination of well-preserved
material collected by Zalessky and myself (1939) proved that they have nothing to do
with Sigillaria. There are no ribs, but Lepidodendron-like leaf -cushions. The leaf-
trace is never visible. These plants are identical with Lepidodendropsis hirmeri
Lutz. Lutz compared Lepidodendropsis with Helenia, but the specimens of this
latter genus, which occurs with Heleniella, are for the most part badly preserved.
Undoubtedly most of the species of Helenia described by Zalessky (1931), especially
H. similis and H. bella, belong to Lepidodendropsis. The same may be true for the
specimens he calls Lepidodendron stylicum, but most of these are very intensively
decorticated.

Zalessky (1930) described a new genus, Micheevia, from the Lower Carboniferous
of the Ural mountains. Some of the species are indeterminable but M. rimnensis,
M. pulchella and M. uralica certainly belong to Lepidodendropsis. M. uralica can
be compared with L. vandergrachti from the Pocono, and M. pulchella with L. hirmeri.

In the same paper he figures Helenia inopinata which also belongs to Lepidoden-
dropsis. The same may be said for his Lepidodendron glincanum (pi. 2, figs. 3, 4),
but the specimens, some of which show their internal structure, are too decorticated
to be certain. Possibly Lepidodendron caracubense Zalessky (1921) ought to be
included in Lepidodendropsis but this is not certain.

It is probable that the specimen described by Schmalhausen (1883) as Lepido-
dendron glincanum Eichw. from the Egorshino region, together with those identified by
Magdefrau (1936, pi. 10, figs. 6-8) as Heleniella theodori from the Upper Devonian
in the Thiiringer Wald, and Gilkinet's Lepidodendron nothum (1922, pi. 13, fig. 76),
all belong to Lepidodendropsis.

Gothan & Zimmermann (1937) described lepidodendroid remains from the Upper
Devonian of Bogendorf-Libichau near Waldenburg. Several of their figures are
very much like Lepidodendropsis but in most cases the specimens are too incomplete
or too badly preserved to be certain. The small stems (pi. 24, figs. 1-4) and the
specimens ascribed to Protolepidodendropsis frickei G. & Z. (pi. 24, figs. 6a, b ; pi. 25,
figs, i, la, ? 7) almost certainly belong to Lepidodendropsis. In my opinion there
is no reason for the creation of a new genus for these stems. Whether they are
specifically identical with any of the species described in Lepidodendropsis is another
question. However, there is a resemblance to L. hirmeri Lutz. Krausel & Weyland
(1949 : 136) also state that this genus is practically indistinguishable from Lepidoden-
dropsis.

Gothan & Zimmermann (1937, pi. 22, figs. 1-4) described and figured a new
species of Sublepidodendron (S. antecedens) from the Upper Devonian of Oberkunzen-
dorf. Krausel & Weyland (1949 : 146) state that it can be compared with Lepidoden-
dropsis. It very probably belongs to this genus.

Hoeg (1942) described a new species of Protolepidodendropsis (P. pulchra Hoeg)
from the Upper Devonian of Mimerdalen, Spitsbergen. He compares it with
Heleniella theodori Zal. from the Donetz Basin, with Protolepidodendron, and with

THE CARBONIFEROUS FLORA OF PERU 219

Protolepidodendropsis frickei. Most of the specimens figured by Hoeg in his pis. 54
and 55 cannot be separated from Lepidodendropsis .

Another group of lepidodendroid plants with which Hoeg compares his new species
is Sublepidodendron, especially 5. subfallax Nath. andS. nordenskioldi Nathorst (1920).
Several of Nathorst's figures resemble Lepidodendropsis at first sight, but the struc-
ture and position of the leaf-cushions in Sublepidodendron do not permit one to unite
them with Lepidodendropsis. The most typical species described by Nathorst from
Spitsbergen, Sublepidodendron mirdbile, S. fallax, S. subfallax and 5. nordenskioldi,
are considered by Gothan (1933) to be synonymous, and he includes them all in one
species Sublepidodendron mirabile Nathorst. Gothan, however, does not use this
generic name and retains the species under Lepidodendron. In my opinion and in
accordance with Nathorst (1920), Hirmer (1927) and Gothan & Zimmermann (1937)
it is necessary to separate them from Lepidodendron. Gothan also unites Lepidoden-
dron leeianum G. & S. with L. mirabile, but Sze (1936) does not agree and considers
the former to be a distinct species of Sublepidodendron.

Possibly Lepidodendron calamitoides Nathorst (1920, pi. 5, figs. 1-8, ? 9) also
belongs to this group. The relationship of L. kidstoni Nathorst (1920, pi. 3, figs.
la, 2-7) is more doubtful. It may be that such specimens are in some respects
already transitional to the true Lepidodendra. There are several species of Lepi-
dodendron in the Namurian and in the upper part of the Mississippian, which resemble
Sublepidodendron in many characters, but their leaf-cushions, and especially their
true leaf -scars, are lepidodendroid. Such species include L. kidstoni Nath. (which
can be compared with Sublepidodendron), L. robertii Nath., L. acuminatum Goepp.,
L. spetsbergense Nath., L. osbornei Walkom and perhaps L. volkmannianum.

Carpentier (1932 : 33) compared the previously described Lepidodendron corrugatum
Dawson (Carpentier, 1925, pi. 13, figs. 6-8) from the Tournaisian of Bois Gamats,
near Laval (Mayenne), with Heleniella theodori Zal. So far as one can judge from his
figures, especially that of 1932, he is quite right in this comparison, and in my opinion
they are at least distantly related to Lepidodendropsis hirmeri. Zalessky (in litt.)
objected to Carpentier 's conclusions and supposed that the French specimens most
probably belonged to a new type. His objections, however, were based on incom-
plete knowledge of his genus Heleniella. A revision of this genus has since been
made, based on new material collected by Zalessky and myself.

Lepidodendropsis is also known from some other localities in Asia. Zalessky
(1937) figures a small poorly preserved specimen from the River Niaysse, 4 km.
from the River Mania-Niaysse, which he names Heleniella theodori. It may belong
to Lepidodendropsis but it is impossible to be certain. From the same locality he
describes Ularia ovalis Zal. which he considers to be a rhizome of a Lepidophyte
(Zalessky, 1937 : 10, pi. 9, fig. i). It is possible that this also is a fragment of
Lepidodendropsis. He compares the specimen with Stigmaria exigua Dawson
(1871, pi. 3, fig. 30, 300) but Dawson's specimen does not allow further determina-
tion.

As already mentioned Sze (1936) does not agree with Gothan's identification of
Lepidodendron leeianum Gothan & Sze with Sublepidodendron mirabile Nath. In
the same paper Sze describes a number of specimens from the Chinese province of

220 THE CARBONIFEROUS FLORA OF PERU

Kiangsu as Lepidodendron aff. leeianum Goth. & Sze (? n. sp.) which very much
resemble Lepidodendropsis. He states that the leaf-scars are not very distinct.
In the figures, especially pi. 2, fig. 2, the leaf-cushions are very similar to those of
Lepidodendropsis, and no leaf-scar is present.

Protolepidodendron (?) arborescens Sze (19360) almost certainly belongs to Lepi-
dodendropsis. It has nothing to do with Protolepidodendron, which is characterized
by leaves divided in their upper parts. Sze (pi. 2, fig. 8) figures a divided leaf, but
there is no proof that it belongs to this species. He compares the species with
P. scharyanum Krejci (Halle 1936, pis. 2, 3 ; text-fig, i) from Yunnan, which, apart
from the division of the leaves, is also much like Lepidodendropsis. I should not
hesitate, if such leaves were absent, to unite it with this genus.

Halle's material shows that at least some of the specimens named Protolepido-
dendron are distinguishable from Lepidodendropsis by the divided leaves only.
Whether the presence or absence of the division in these leaves is sufficient ground
for a generic separation (cf. Krausel & Weyland 1949 : 136) is, in my opinion,
rather doubtful. Sze also doubts whether such specimens as those figured by
Halle belong to Protolepidodendron.

Walkom (1928) described two new species of Protolepidodendron (P. linear e and
P. yalwalense} from Yalwal, New South Wales. He compared the first with P.
primaevum (White) which has since been named Lepidosigillaria (Krausel & Weyland,
1949 : 148), and P. yalwalense with Lepidodendron karakubense Schmalh. which is
possibly Lepidodendropsis. Both the plants used for comparison by Walkom have
been recorded from beds considered to be Upper Devonian, and for this reason
Walkom also includes the Yalwal beds in the Upper Devonian, although they are
considered to be Carboniferous by Clarke and other authors.

At a number of localities the flora, as far as it is known, does not contain Lepidoden-
dropsis, but Rhacopteris of the ovata group only. Such is the case in Australia
(Feistmantel 1890), some parts of South America, and also in Spiti, India, the flora
of which has been described by Gothan & Sahni (1937).

It is clear that the Lepidodendropsis (Rhacopteris} flora, or at least related floras,
are found all over the world, and this is very important for the stratigraphy of the
Lower Carboniferous and for comparison with the Upper Devonian. It is interest-
ing to note that such plants, especially Lepidodendropsis, also occur in different
localities which are considered as Upper Devonian. This fact is not surprising,
since in many cases it is almost impossible to separate what is considered to be
Upper Devonian from the Lower Carboniferous. In some cases these so-called
Upper Devonian floras are typical Lepidodendropsis floras, as in the Donetz Basin.

It is possible that the Lepidodendropsis-Rhacopteris floras represent a transition
between the Archaeopteris-Cyclostigma floras of the Devonian and the floras of the
lower part of the Mississippian.

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PRESENTED

5 OCT 1954

EXPLANATION OF PLATES

The specimens figured in PI. 17-20 are in the Department of Geology, British Museum (Natural
History). Those figured in Pis. 21-26 belong to the Geologisch Bureau, Heerlen. All the
photographs were taken by Mr. Van Voskuijlen, Geologisch Bureau, Heerlen.

Unless otherwise stated all the figures are natural size.

PLATE 17

Sphenopteris whitei (Berry) and Cyclostigma pacifica (Steinmann).
Lower Carboniferous (Mississippian) ; Paracas, Peru.

FIG. i. Sphenopteris whitei (Berry). ¥.25936.

FIG. 2. Sphenopteris whitei (Berry). Counterpart of fig. i. ¥.25936(3.

FIG. 3. Sphenopteris whitei (Berry). V. 25942.

FIG. 4. Sphenopteris whitei (Berry). Part of counterpart of fig. i. V. 259366.

FIG. 4<* Enlargement of fig. 4, x 3.

FIG. 5. Cyclostigma pacifica (Steinmann). V. 25925.

FIG. 5» Enlargement of fig. 5, x 3.

FIG. 6. Cyclostigma pacifica (Steinmann). V.25925a.

FIG. 6a Enlargement of fig. 6, x 3.

FIG. 7. Cyclostigma pacifica (Steinmann). V. 25928.

Bull. B.M. (N.H.) Geol. 2, 5

PLATE 17

Photo's van Voskuylen

SPHENOPTERIS, CYCLOSTIGMA

PLATE 18

Cyclostigma pacifica (Steinmann).
Lower Carboniferous (Mississippian) ; Paracas, Peru.

FIG. 8. Cyclostigma pacifica (Steinmann). ¥.25920.

FIG. 8a Enlargement of fig. 8, x 3.

FIG. 9. Cyclostigma pacifica (Steinmann). V. 25918.

FIG. ga Enlargement of fig. 9, x 3.

FIG. 10. Cyclostigma pacifica (Steinmann). V.259i8a.

FIG. loa Enlargement of fig. 10, x 3.

Bull. B.M. (N.H.) Geol. 2, 5

PLATE 18

CYCLOSTIGMA

PLATE 19

Cyclostigma pacifica (Steinmann).
Lower Carboniferous (Mississippian) ; Paracas, Peru.

FIG. ii. Cyclostigma pacifica (Steinmann). ¥.25917.

FIG. 12. Cyclostigma pacifica var. decor ata Jongmans. ¥.25948.

FIG. i2a Enlargement of fig. 12, x 3.

FIG. 13. Cyclostigma pacifica (Steinmann). ¥.25933.

FIG. 14. Cyclostigma pacifica var. decorata Jongmans. ¥.25919.

FIG. i4« Enlargement of fig. 14, x 3.

FIG. 14^ Cyclostigma pacifica (Steinmann). ¥.25916. Enlargements (figs. 146, and 146,)
on PI. 20.

Bull. B.M. (N.H.) Geol. 2, 5

PLATE 19

14bi

14 a

CYCLOSTIGMA

PLATE 20

Cyclostigma pacifica (Steinmann), Lepidodendropsis peruviana (Gothan),
? Lepidodendropsis lissom (Steinmann), Lepidodendropsis sp., Rhacopteris cf. cuneata
(Walkom) and Rhacopteris ovata (McCoy).

Lower Carboniferous (Mississippian) ; Paracas, Peru.

FIG. 1462 Cyclostigma pacifica (Steinmann). Enlargement of fig. i^j, x 3.

FIG. 1463 Cyclostigma pacifica (Steinmann). x 3.

FIG. 15. Lepidodendropsis peruviana (Gothan). V. 25911.

FIG. 16. ? Lepidodendropsis (Lepidodendron] lissoni (Steinmann). ¥.25932.

FIG. 17. ? Lepidodendropsis sp. ¥.25915.

FIG. 18. Rhacopteris cf. cuneata (Walkom). V. 25948*1.

FIG. i8a Rhacopteris cf. cuneata (Walkom) and Rhacopteris ovata (McCoy). The specimen

is on the reverse side of that shown in fig. 12.
FIG. T.8al Rhacopteris cf. cuneata (Walkom). Enlargement of fig. i8a, x 3.

Bull. B.M. (N.H.) Ceol. 2, 5

PLATE 20

16

15

CYCLOSTIGMA, LEPIDODENDROPSIS. RHACOPTFRTS

PLATE 21

Lepidodendropsis de voogdi n. sp.
Lower Carboniferous (Mississippian) ; Carhuamayo, Peru.

FIG. 19. Lepidodendropsis de voogdi Jongmans. Photo 8414.

FIG. 20. Lepidodendropsis de voogdi Jongmans. Traces of the sporangia are visible on the

left of the figure. Photo 8416.

FIG. 21. Lepidodendropsis de voogdi Jongmans. Photo 8444.
FIG. 2ia Enlargement of fig. 21, x 3.
FIG. 22. Lepidodendropsis de voogdi Jongmans, showing sporangia containing megaspores.

Photo 8413.

FIG. 22<* Enlargement of fig. 22, x 3.
FIG. 23. Lepidodendropsis de voogdi Jongmans. Photo

Bull. B.M. (N.H.) Geol. 2, 5

PLATE 21

LEPIDODENDROPSIS

PLATE 22

Lepidodendropsis de voogdi n. sp., and Lepidodendropsis steinmanni n. sp.
Lower Carboniferous (Mississippian) ; Carhuamayo, Peru.

FIG. 24. Lepidodendropsis de voogdi Jongmans. Photo 8445.

FIG. 25. Lepidodendropsis de voogdi Jongmans. Photo 8449.

FIG. 26. Lepidodendropsis de voogdi Jongmans. Photo 8443.

FIG. 27. Lepidodendropsis de voogdi Jongmans. Photo 8448.

FIG. 28a Lepidodendropsis steinmanni Jongmans, showing habit of the stem with leaves.

Photo 8415.
FIGS, z&d, z8da. Pith-cast, belonging to specimens in figs. 280 and 28b (PI. 23). Photo 8450.

Fig. zSda, x 3.

Bull. B.M. (N.H.) Geol. 2, 5

PLATE 22

LEPIDODENDROPSIS

PLATE 23

Lepidodendropsis steinmanni n. sp. and Cyclostigma cf. pacifica (Steinmann) var.
Lower Carboniferous (Mississippian) ; Carhuamayo, Peru.

FIG. 286 Lepidodendropsis steinmanni Jongmans. Counterpart of fig. 28a with better

preserved leaves. Photo 8415.

FIG. 28c Enlargement of part of Fig. 286 showing the leaves, x 3. Photo 8415.
FIG. 29a Cyclostigma cf. pacifica (Steinmann) var. Upper surface of stem. Photo 8451.
FIG. zgaa Enlargement of fig. 290, x 3.

Bull. B.M. (N.H.) Geol. 2, 5

PLATE 23

T.FPTDOnRNDTCOPSIS. CYCLOSTIGM A

GEOL. II, 5.

PLATE 24

Cyclostigma pacifica (Steinmann) and Lepidodendropsis cf. steinmanni Jongmans.
Lower Carboniferous (Mississippian) ; Carhuamayo, Peru.

FIG. 2gb Cyclostigma cf. pacifica (Steinmann) var. Lower surface of stem. Photo 8451.

FIG. 2gba Enlargement of fig. 296, x 3.

FIG. 2gc Cyclostigma cf. pacifica (Steinmann) var. Pith-cast belonging to figs, zga and

296. Photo 8451.

FIG. 2Qca Enlargement of fig. igc, x 3.

FIG. 30. Cyclostigma pacifica (Steinmann). With leaves attached. Photo 8532.
FIG. 300 Enlargement of fig. 30, x 3.
FIG. 31. Lepidodendropsis cf. steinmanni Jongmans. Photo 8447.

Bull. B.M. (N.H.) Geol. 2, 5

PLATE 24

CYCLOSTIGMA, LEPIDODENDROPSIS

PLATE 25

Lepidodendropsis cf . de voogdi Jongmans, Rhacopteris ovata (McCoy)
and Rhacopteris cf. circularis Walton.

Lower Carboniferous (Mississippian) ; Carhuamayo, Peru.

FIG. 32. ? Lepidodendropsis cf. de voogdi Jongmans. Photo 8446.

FIG. 33. Rhacopteris ovata (McCoy). Photo 8456^.

FIG. 34. Rhacopteris ovata (McCoy). Photo 84566.

FIG. 35. Rhacopteris ovata (McCoy). Photo 8452.

FIG. 36. Rhacopteris ovata (McCoy). Photo 8454.

FIG. 37. Rhacopteris ovata (McCoy). Photo 8453.

FIG. 38. Rhacopteris cf. circularis Walton. Photo 84560.

Bull. B.M. (N.H.) Geol. 2, 5

PLATE 25

32

34

LEPIDODENDROPSIS, RHACOPTERIS

PLATE 26

Rhacopteris cf. circularis Walton, Rhacopteris ovata (McCoy), Triphyllopteris collom-
biana (Sch.), Triphyllopteris lescuriana (Meek) and ? Triphyllopteris peruvian*
Jongmans.

Lower Carboniferous (Mississippian) ; Carhuamayo, Peru.

FIG. 39. Rhacopteris cf. circularis Walton. Photo 84560.

FIG. 40. Triphyllopteris collombiana (Sch.) Photo 8455.

FIG. 41. Triphyllopteris collombiana (Sch.) Photo 8455^

FIG. 42. Triphyllopteris collombiana (Sch.) Photo 84556.

FIG. 43. Triphyllopteris lescuriana (Meek). Photo 845801.

FIG. 44. Triphyllopteris lescuriana (Meek). Photo 84586.

FIG. 45. Triphyllopteris lescuriana (Meek) and Rhacopteris ovata (McCoy). Photo 8458.

FIG. 46. ? Triphyllopteris peruviana Jongmans. Photo 8457.

Bull. D.M. (N.H.) Geol. 2, 5

PLATE 26

RHACOPTERIS, TRIPHYLLOPTERIS

PRESENTED
5 OCT 1954

PRINTED IN GREAT BRITAIN BY
ADLARD AND SON, LIMITED
BARTHOLOMEW PRESS, DORKING

FURTHER CONTRIBUTIONS

TO THE SOLUTION OF THE

PILTDOWN PROBLEM

J. S. WEINER; W. E. LE GROS CLARK; K. P. OAKLEY;

G. F. CLARINGBULL & M. H. HEY;

F. H. EDMUNDS; S. H. U. BOWIE & C. F. DAVIDSON;

C. F M. FRYD; A. D. BAYNES-COPE;

A. E. A. WERNER & R. J. PLESTERS

BULLETIN OF
THE BRITISH MUSEUM (NATURAL HISTORY)

GEOLOGY Vol. 2 No. 6

LONDON: 1955

BULLETIN OF
THE BRITISH MUSEUM (NATURAL HISTORY)

GEOLOGY

The following papers appeared in Volume I (1949-52) :

Price
No. i (1949). The Pterobranch Rhabdopleura in the English Eocene.

H. D. Thomas & A. G. Davis 75. 6d.

No. 2 (1949)' A Reconsideration of the Galley Hill Skeleton. K. P.

Oakley & M. F. Ashley Montagu ...... 55.

No. 3 (1950). The Vertebrate Faunas of the Lower Old Red Sandstone

of the Welsh Borders. E. I. White.

Pteraspis leathensis White a Dittonian Zone-Fossil. E. I.

White 75. 6d.

No. 4 (1950). A New Tithonian Ammonoid Fauna from Kurdistan,

Northern Iraq. L. F. Spath ....... IDS.

No. 5 (1951). Cretaceous and Eocene Peduncles of the Cirripede Euscal-

pellum. T. H. Withers ........ 55.

No. 6 (1951). Some Jurassic and Cretaceous Crabs (Prosoponidae) .

T. H. Withers 5s.

No. 7 (1952). A New Trochiliscus (Charophyta) from the Downtonian

of Podolia. W. N. Croft IDS.

No. 8 (1952). Cretaceous and Tertiary Foraminifera from the Middle

East. T. F. Grimsdale los.

No. 9 (1952). Australian Arthrodires. E. I. White .... 155.
No. 10 (1952). Cyclopygid Trilobites from Girvan. W. F. Whittard . 6s.

FURTHER CONTRIBUTIONS TO THE
SOLUTION OF THE PILTDOWN PROBLEM

BY

J. S. WEINER; W. E. LE GROS CLARK; K. P. OAKLEY;

G. F. CLARINGBULL & M. H. HEY;

F. H. EDMUNDS; S. H. U. BOWIE & C. F. DAVIDSON;

C. F. M. FRYD; A. D. BAYNES-COPE;

A. E. A. WERNER & R. J. PLESTERS

Pp. 225-287 ; Pis. 27-31

BULLETIN OF
THE BRITISH MUSEUM (NATURAL HISTORY)

GEOLOGY Vol. 2 No. 6

LONDON : 1955

THE BULLETIN OF THE BRITISH MUSEUM
(NATURAL HISTORY) instituted in 1949, is
issued in five series corresponding to the Departments
of the Museum, and an Historical Series.

Parts will appear at irregular intervals as they become
ready. Volumes will contain about three or four
hundred pages, and will not necessarily be completed
within one calendar year.

This part is Vol. 2, No. 6 of the Geological series.

Issued 2ist January, 1955 Price One Pound

FURTHER CONTRIBUTIONS TO THE
SOLUTION OF THE PILTDOWN PROBLEM

CONTENTS

Page
INTRODUCTION. Sir GAVIN DE BEER, F.R.S. ...... 228

1. Outline of the Piltdown Problem. J. S. WEINER .... 229

2. An Anatomical Study of the Piltdown Teeth and the so-called Turbinal

Bone. W. E. LE GROS CLARK, F.R.S. ...... 234

3. The Piltdown " Implements." K. P. OAKLEY ..... 243

4. The Piltdown Mammalia. K. P. OAKLEY ..... 247

5. The Composition of the Piltdown Hominoid Remains. K. P. OAKLEY 254

6. Chemical Changes in Bones : a Note on the Analyses. C. F. M. FRYD 266

7. The X-ray Crystallography of the Piltdown Fossils. G. F. CLARINGBULL

& M. H. HEY 268

8. The Black Coating on the Piltdown Canine. A. E. A. WERNER & R. J.

PLESTERS ........... 271

9. The Geology of the Piltdown Neighbourhood. F. H. EDMUNDS . . 273

10. The Radioactivity of the Piltdown Fossils. S. H. U. BOWIE & C. F.

DAVIDSON ........... 276

11. The Fluorimetric Determination of Uranium in the Piltdown Fossils.

A. D. BAYNES-COPE ......... 283

12. References ........... 285

TABLES :

I. Composition of Stains on Piltdown Flints .... 245

II. Radioactivity of Piltdown Elephant teeth and other Villa-

franchian Fossils ......... 249

III-IV. Analyses of Piltdown Hominoid Bones and Teeth . . . 262
V. Analyses of Piltdown Mammalian Bones and Teeth . . . 263
VI-VII. Analyses of Bones and Teeth used in comparison . . . 264-265
VIII-IX. X-ray Examination of Piltdown Specimens and of Bones Arti-
ficially Iron-stained ........ 269-270

X-XI. Radiometric Assays of Piltdown Specimens and various Tertiary,

Pleistocene and Holocene fossils ...... 279-282

XII. Uranium Content of Piltdown and other Fossils . . . 284

PLATES :

27. Piltdown Mandible and Orang Mandible compared

28. Piltdown Flint " Implements "

29. Piltdown Bone "Implement"

30. Electron-micrograph of Piltdown Mandible and Autoradio-

graph of Piltdown Elephant Molar

31. X-ray Diffraction Photographs of Apatite and Gypsum in

Piltdown and other Bones

GEOL. II, 6. ig.

228 THE SOLUTION OF THE PILTDOWN PROBLEM

INTRODUCTION

WE are now in a position to give an account of the full extent of the Piltdown hoax.
The mandible has been shown by further anatomical and X-ray evidence to be almost
certainly that of an immature orang-utan ; that it is entirely Recent has been con-
firmed by a number of microchemical tests, as well as by the electron-microscope
demonstration of organic (collagen) fibres ; the black coating on the canine tooth,
originally assumed to be an iron encrustation, is a paint (probably Vandyke brown) ;
the so-called turbinal bone is shown by its texture not to be a turbinal bone at all,
but thin fragments of probably non-human limb-bone ; all the associated flint
implements have been artificially iron-stained ; the bone implement was shaped by
a steel knife ; the whole of the associated fauna must have been " planted," and it
is concluded from radioactivity tests and fluorine analysis that some of the specimens
are of foreign origin. The human skull fragments and some of the fossil animal bones
are partly replaced by gypsum, the result of their treatment with iron sulphate to
produce a colour matching that of the gravel. Not one of the Piltdown finds genuinely
came from Piltdown. These latest investigations have demonstrated the methods
now available which will not only make a successful repetition of a similar type of
forgery virtually impossible in the future, but will be of further value in palaeonto-
logical research.

GAVIN DE BEER,

Director.

i. OUTLINE OF THE PILTDOWN PROBLEM

By J. S. WEINER

Department of Anatomy, University of Oxford

THERE are three sites on or near Piltdown Common, Sussex, to which attention was
originally drawn by Charles Dawson and where in the years 1908 to 1915 fossil and
archaeological remains came to light. The first site is a gravel pit alongside the main
drive in the grounds of Barkham Manor, about a hundred yards from the house.
Here fossils and implements were found at intervals from 1908 to 1914. Smith
Woodward was first informed of the skull (here referred to as Piltdown I) in a letter
from Dawson dated I4th February, 1912. The gravel itself was noticed by Dawson
as a likely source of Pliocene or Early Pleistocene fossils some years earlier, possibly
in 1899 (Dawson, 1913 : 75). The second site is a field (its precise situation is not
recorded) probably at Sheffield Park about two miles to the north-west of the first
site. Here finds (Piltdown II) were reported by Dawson in 1915. The third site is
at Barcombe Mills, four miles south-west of Piltdown ; cranial fragments said to
have been obtained here were in Dawson's possession at the time of his death and
were obtained for the British Museum by Smith Woodward. Neither the date of
this discovery nor the exact spot has been recorded. Only at Barkham Manor were
any systematic excavations carried out ; these were conducted by Smith Woodward
and Dawson in 1912, 1913 and 1914, by Woodward alone in 1916 — the year of
Dawson's death — and spasmodically afterwards (Woodward, 1948). Nothing was
found in the 1916 season, nor when gravel was dug at another spot nearer the farm-
yard some years later. The original site was re-excavated in 1950 under the auspices
of the Nature Conservancy and a new trench dug to provide a permanent witness
section of the Piltdown gravel (Toombs, 1952), but nothing human or animal came
to light. The material from all three sites is considered in this report.

The bibliography on Piltdown is very large ; over 300 references were listed by
W. & A. Quenstedt (1936), and a select list was given by Oakley (1952). The teeth
and the cranial and mandibular fragments (Piltdown I and II) were described by
Woodward (Dawson & Woodward, 1913, 1914 ; Woodward, 1917), and by Keith
(1925), and the Barcombe Mills skull fragments by Ashley Montagu (1951). Accounts
of the animal remains have been given by Woodward (Dawson & Woodward, 1914,
1915) and by Hopwood (1935). The flint implements were dealt with by Dawson in
the first publications and by Woodward in The Earliest Englishman (1948). The
unique bone implement was the subject of a special paper (Dawson & Woodward,

The circumstances and sequence of the finds, which have an important bearing on
the controversies arising from Smith Woodward's interpretation and on the final
invalidation of his conclusions by the disclosure of the fraudulent nature of the

230 THE SOLUTION OF THE PILTDOWN PROBLEM

material, will be dealt with by one of us (J. S. W.) in a forthcoming book to be
published by the Oxford University Press.

Smith Woodward accorded the finds at Barkham Manor full recognition as a new
genus and species of the Hominidae — Eoanthropus dawsoni— oi Early Pleistocene
age, " defined by its ape-like mandibular symphysis, parallel molar-premolar series,
and narrow lower molars which do not decrease in size backwards ; to which diag-
nostic characters may probably be added the steep frontal eminence and slight
development of brow-ridges". Both the dating and the attribution of the remains
to one individual were challenged at the first announcement of the discovery on
i8th December, 1912. Newton and Keith favoured a Pliocene date (Dawson &
Woodward, 1913), but the majority agreed with Dawson's and Woodward's Lower
Pleistocene estimate, which remained undisputed until Edmunds, in 1925, made a
new geological survey of the East Sussex region. In 1935 Hopwood placed the skull
and jaw with the derived Villafranchian remains. The anatomical conclusion was
disputed strongly by Waterston from the start (in Dawson & Woodward, 1913 : 150 ;
Waterston, 1913) and by Miller (1915), Boule (1915), and others. To these authors
the combination of a skull and mandible of such distinct types in a single individual
seemed, on morphological grounds, highly improbable. They pointed out that the
skull bones, if found by themselves, would certainly have been referred to the genus
Homo, and the mandible, if found by itself, would as certainly have been accepted
as that of an anthropoid ape. Thus two distinct forms were presumed to be present
— fossil Homo sapiens, and a fossil ape named Pan vetus by Miller (1915) and Boreo-
pithecus dawsoni by Friederichs (1932).

None the less, the evidence and arguments put forward by Woodward made a
coherent and convincing case which was enormously strengthened by the finding of
the canine in 1913 and Piltdown II in 1915. Keith restated and extended this case
in 1915, giving his support to Woodward's thesis after careful weighing of the
evidence. Not for many years could the evidence in any respect, anatomical, geo-
logical, archaeological or phyletic be shown with certainty to be untenable, though
a number of serious criticisms were brought forward. On the available evidence
the strength of the case for E. dawsoni may be judged by listing the arguments
urged in favour of Woodward's interpretation:

1. The probable natural association of cranium with jaw was attested by
their close physical proximity, especially so in a gravel formation — the mandible
" within a yard " of where one piece of occiput had been found by Woodward
in an undisturbed patch of gravel (Dawson, 1913), the canine in gravel " in
situ excavated within a radius of 5 yards of the spot where the mandible was
found " ; the nasal bones and " turbinal " within 2 or 3 ft. of the mandible.

2. The complementary nature of the fragments — mandible, canine and jawless
cranium — in such close physical proximity pointed irresistibly to their natural
association.

3. The state of mineralization, colour, and unrolled condition of the different
pieces appeared very similar. The canine tooth, like the bony fragments, was
apparently iron-stained.

4. There was positive anatomical evidence of the natural association between

THE SOLUTION OF THE PILTDOWN PROBLEM 231

lower jaw and cranium. In particular, the remarkable flat wear on the molars
was a human and not an ape-like character, and functionally in entire accord
with the human type of articulation of the glenoid cavity.

5. The canine in its wear also indicated a complete departure from the
normal ape-like condition. The canine could not have overlapped in the normal
simian fashion with the corresponding tooth in the opposing jaw since there is
no attrition facet on either the proximal or distal aspect of the tooth.

6. The X-ray appearance of the roots of the molar tooth was much more
reminiscent of the human than the ape condition (Keith, 1925).

7. Other arguments, of less weight than the foregoing, were (i) Pycraft's
(1917) belief that the axis of alignment of the molar teeth in the jaw was much
more like man than ape, (ii) Elliot Smith's view that the endocranial cast
showed simian features and (iii) Woodward's (1932) inference that the order of
eruption was human and not ape-like based on the much greater attrition
suffered by the canine in comparison with that of the molar.

8. As a further refutation of the idea that the association of jaw and cranium
was an accidental coincidence, there was the discovery of Piltdown II. The
finding of another molar of the same type as those of Piltdown I, also associated
with cranial fragments denoting a second individual not distinguishable from
Homo, increased very greatly the probability of the natural association of
mandible and brain-case.

9. The combination of hominid skull with ape-like jaw was not inadmissable
• on grounds of morphological incompatibility since there are many fossil instances

of quite unexpected combinations of skeletal structures.

10. The existence of a creature like E. dawsoni was consistent with its reported
geological age, as judged by associated fossils and the apparently " pre-Chel-
lean " tools, for an ancestral or transitional form of this kind was to be expected
in the Lower Pleistocene.

11. " Piltdown Man " was more simian than Heidelberg Man. Though
morphologically very different from Pithecanthropus (then known only by skull
cap, two doubtful molars and a disputed femur) it had as good a claim to
represent the ancestor of H. sapiens since in brain size it was far more advanced
than Pithecanthropus.

Woodward's interpretation was thus a close-knit set of arguments which took all
the evidence into account. The alternative hypothesis that the discoveries repre-
sented two distinct creatures — fossil man and fossil ape — could not account for all the
evidence without raising new complexities. To avoid the acceptance of Piltdown II
as a second remarkable coincidence some doubted its authenticity. Hrdlicka (1922)
suggested that the isolated molar must have come from the first site and that some
mistake had been made — a suggestion denied by Woodward (1933). Weinert (1933)
thought that the frontal bone of Piltdown II really belonged to Piltdown I. Weiden-
reich (1937) supposed that the isolated molar was human, so making Piltdown II a
discovery of prehistoric H. sapiens only.

An anatomical argument against the association of the cranium with the mandible

232 THE SOLUTION OF THE PILTDOWN PROBLEM

is that the bicondylar width of the mandible does not correspond to the distance
between the mandibular fossae on the base of the skull, and therefore it is impossible
to fit the mandible to the skull. But the symphysial region of the mandibular
fragment is missing, and with no certain evidence of the position of the mid-line
of the mandible there can be no certainty in estimating the bicondylar width.

However much evidence of the ape-like character of the mandible was brought
forward (Miller, Ramstrom, Friederichs), the inference that the missing crucial
condylar region would also be ape-like remained incapable of proof. With so much
else ape-like, a variety of suggestions were made to account for the " un-ape-like "
dental wear. Miller (1915) thought that similar flat wear might occasionally occur
even in modern apes. The specimen which he adduced, however, was not only
unusual but quite abnormal (Pycraft, 1917). Weidenreich (1937) drew attention to
flat wear in the molars of a Pleistocene orang in Mme. Selenka's collection from Java
but he only reproduced a photograph of this specimen and made no detailed com-
parison with the Piltdown molars. Both he and Miller left the peculiar wear on the
canine unexplained. Marston (1952) has attempted to go further and has theorised
as to the movements which might produce the Piltdown wear, supposing that the
canine were an upper canine. But that such movements were ever made by a jaw
with a structure, so far as it is known, and muscular attachments indistinguishable
from that of modern apes remains completely hypothetical.

Nevertheless, certain of the criticisms of Woodward's conclusions cannot be
disregarded. In particular, the " human " features said to exist in the mandible
and teeth are few indeed (though they are crucial), whereas detailed study serves
only to emphasise the astonishing similarity of the mandible to that of a modern
orang or chimpanzee. Sicher's study (1937) deserves mention here, for, impressed
by the completely non-human configuration of the dental foramen and its rela-
tion to the mandibular canal, Sicher questioned the association of the jaw and
cranium. Keith (1925) himself did much to throw doubt on certain alleged simian
features of the braincase and endocranial cast brought forward by Elliot Smith.
Symington (1915) severely criticized the latter's conclusions, and could find no
convincing evidence of any precocious or peculiar development in the brain of
Eoanthropus. Lyne (1916) drew attention to the extraordinary contradiction be-
tween the apparent immaturity of the canine and its excessive wear, but his explana-
tion that the tooth might be a milk canine had little to support it and his important
observations were disregarded. Miller (1918) and Marston (1952) threw doubt on
Pycraft's belief in the " near-human " alignment of the molars in the jaw. Doubts
had early been expressed on the workmanship of the bone implement (Reginald
Smith, in discussion of Dawson & Woodward, 1915 ; Breuil, 1938) ; these were
renewed by Oakley in 1949.

The interpretations of Woodward and his critics were beset with new difficulties
when fluorine tests (Oakley & Hoskins, 1950) showed that E. dawsoni could not be
accepted as a Late Pliocene derivative, but was apparently contemporary with the
gravel. Combined with Edmunds' conclusion of 1926, this implied the dating of the
remains to the last interglacial period, i.e. the early part of the Upper Pleistocene.
These difficulties were further enhanced from about 1925 onwards by many new

THE SOLUTION OF THE PILTDOWN PROBLEM 233

discoveries of Pithecanthropus (both in Java and China), of the Australopithecinae
and of fossil apes (particularly in E. Africa.)

By 1950 every possible opinion of " Piltdown Man's " status had been discussed.
Weidenreich (1947) had decided to dismiss the " chimaera " altogether ; Friederichs
(1932), Montagu (1951), Marston (1952) and others believed that the remains repre-
sented two distinct fossil creatures ; some (Howells, 1947 ; Leakey, 1953) felt that
the situation was so confused that no definite decision could be made ; others, that
the matter should be left in " suspense account " (Le Gros Clark, 1949) in the hope
that more material might be found ; and there were some for whom " Eoanthropus
dawsoni " continued to figure unquestioned as a species or genus of the Hominidae.
Finally, Weinert (1953) thought that the jaw, if properly reconstructed, would turn
out to be hominid and by no means ape-like.

The hypothesis of a fake or hoax

In the new situation created by the revised dating the only two conceivable
" natural " explanations both seemed entirely inadequate on evolutionary grounds
and the many puzzling anatomical features remained unresolved. How else could
all the apparent facts be explained ?

The possibility had to be faced that the Piltdown finds were a hoax — that the
mandible was indeed that of an ape, but of a modern ape so treated by mutilation
of the fragment, abrasion of the teeth and staining as to appear a genuine fossil.
Not only could this hypothesis at once explain the circumstances of the finds and
the sequence of discoveries, it could also be entertained on the following grounds:

(1) That the only morphological feature in the mandible which could not
be said to be ape-like was the wear on the molars and it seemed surprising that
this should be the only undoubted feature to link the cranium and mandible.

(2) That filing down of chimpanzee and orang molars was found to produce an
appearance similar to that of the Piltdown molars.

(3) That artificial abrasion would explain the baffling and unique wear of
the canine.

(4) That the very parts of the mandible which one would expect a faker to
remove were, in fact, broken off.

(5) That Oakley (in 1949) found the dentine under the thin, dark " ferru-
ginous" layer to be "most unexpectedly — pure white". (Oakley & Hoskins,
1950:381).

(6) That the 1949 fluorine analysis had left the antiquity of the mandible
quite indeterminate. It had, in fact, " failed to differentiate Eoanthropus from
Holocene bones". (Oakley, 1951 : 50).

(7) That an element of doubt already surrounded the bone implement.

An exhaustive re-examination of all the Piltdown finds has completely confirmed
the hypothesis of a hoax, and experimental work has shown that all the features
of the Piltdown teeth and jawbone can be reproduced artificially. The main results
have already been published (Weiner, Oakley & Le Gros Clark, 1953). The present
series of reports gives the evidence in greater detail and covers a wider field.

2. AN ANATOMICAL STUDY OF THE

PILTDOWN TEETH AND THE SO-CALLED

TURBINAL BONE

By W. E. LE GROS CLARK, F.R.S.

Department of Anatomy, University of Oxford
(I) THE ANATOMICAL DETAILS OF THE PILTDOWN TEETH

THE teeth from Piltdown are the first and second right lower molars in the mandibular
fragment, an isolated canine tooth, and an isolated left lower molar tooth. The last
was reported to have been found in a heap of stones raked off a ploughed field about
two miles from the original Piltdown site. Considered by themselves, it seems
certain that the teeth would have been attributed to an anthropoid ape (quite
similar to a chimpanzee or an orang) except for two main features : (i) the extremely
flat wear of the molar teeth in the mandible, which is not normally to be found in
pongid teeth at an equivalent stage of attrition, but which closely approximates to
the type of wear commonly found in hominid molars ; and (2) the quite unusual
type of wear on the canine which, so far as we are aware, is not paralleled in the
canines of any of the known genera of anthropoid apes, recent or extinct. The
suggestion that these aberrant features might be the result of artificial abrasion at
once offered a plausible explanation of their seemingly anomalous character. Indeed,
it may well be asked why such a suggestion had not been seriously considered until
quite recently. There are no doubt several reasons for this. In the first place, the
mandibular fragment and the canine tooth were reported to have been found by
experienced palaeontologists during their excavations at Piltdown, and the occurrence
in situ has thus always been accepted without question. Secondly, the faking,
obvious though it now appears, had been accomplished with extraordinary skill ;
and, lastly, the statement that the worn surface of the canine shows an exposure of
secondary dentine would almost certainly have distracted attention from a possible
consideration of faking by artificial abrasion. Secondary dentine is deposited as a
reaction to prolonged natural wear and its presence in the canine would thus pre-
suppose that the excessive wear of this tooth was indeed natural. In fact, a re-
examination of the canine has shown that there is no evidence of the deposition of
secondary dentine.

Two other relevant features have been held by some authorities to distinguish
the Piltdown molars of the mandible from those of anthropoid apes. One is their
hypsodont character. But comparative study has shown that, while a similar degree

THE SOLUTION OF THE PILTDOWN PROBLEM

235

of hypsodonty is very unusual (if, indeed, it does occur) in chimpanzees and gorillas,
it is not uncommon in the orang. The other feature is the relative shortness of the
roots of the molar teeth as they appeared to be displayed in the original radiograph
of the Piltdown molars published in a paper by Underwood (1913) and subsequently
copied in publications by other authors (e.g. Keith, 1915 ; Lyne, 1916). But new
radiographs taken recently show quite clearly that the roots are in fact markedly
longer than they have been portrayed, and are thus entirely simian in appearance.
This seems to us to be an important point which needs to be emphasized. The original
radiographs lacked sufficient definition to outline the roots distinctly : from the
recently taken radiograph the outlines of the molars have been reproduced for
comparison with those made from appearances shown in the original radiograph
(Text-figs, i, 2). It will be observed that the lower end of the anterior root of Mi

TEXT-FIG, i . A. Outline drawing of the molar teeth in the Piltdown mandible showing the
roots as they had been interpreted on the basis of the original radiograph published
in 1913. B. A similar drawing made from a recent radiograph showing the actual
form and extent of the roots. The anterior root of Mi has been broken off, and its
probable extent is indicated by a broken line. In both figures the position of the
mandibular canal is shown. Natural size.

TEXT-FIG. 2. Radiograph of the two molar teeth in the Piltdown mandible showing
the form and extent of their roots. Note the apparently accurate apposition of the
crowns of the teeth at their contact facets. Twice natural size. [X-ray by P. E.
Purves.]

236 THE SOLUTION OF THE PILTDOWN PROBLEM

has been broken off (presumably it has been involved in the fracture of the mandible
at this level). Its total length can therefore not be precisely estimated, but there is
an indication of the lower end of the socket which suggests that originally the apex
of the root curved downwards and backwards for some distance. The posterior root
of Mi is long and is deflected backwards at its lower extremity. The radiograph of
the mandible shows that it actually reaches the upper border of the mandibular canal.
The anterior root of M.2 is well defined and was correctly displayed in the original
radiograph. Compared with the posterior root of Mi, the rounded and blunt apex
of this root and the relative width of the apical canal suggest that the root had per-
haps not completed its full development. The posterior root of M2 is considerably
longer than the anterior root, extending downwards to the level of the lower border
of the mandibular canal. This relationship is of some significance, for the " hominid "
appearance in the original radiograph showing the roots of both the first and second
molars apparently falling well short of the canal certainly misled some authorities.
Like the anterior root, the apex of the posterior root of M2 is bluntly rounded and
the apical canal relatively wide.

The problem having been posed — is the unusual type of wear of the Piltdown
molar teeth the result of natural attrition during life or of artificial abrasion after
death ? — consideration was given to those details which on close inspection might
be expected to differentiate the one from the other. A critical study of the teeth
at once revealed certain features which had either escaped notice previously, or the
possible significance of which had not been realized. Indeed, it was because these
features appeared to lend such strong support to the hypothesis of artificial abrasion
that it was decided to re-examine all the Piltdown material for further evidence of
faking.

The flatness of the molar teeth is astonishingly even over almost the entire extent
of the occlusal surface, as though, indeed, the latter had been planed down by some
rapidly acting shearing force. A considerable area of dentine (about 4 mm. in its
greatest diameter) has been exposed on the antero-internal cusps of both teeth, and
not only are these areas quite flat, they are also flush with the surrounding enamel
(Text-fig. 3, a, b). But in natural attrition, whether in hominids or pongids, areas

_....

abed

TEXT-FIG. 3. Diagram illustrating the peculiar type of abrasion on the cusps of the
Piltdown molar teeth. In (a) is shown a schematic section through an unworn cusp.
If this were subjected to artificial abrasion in the plane indicated, the appearance
shown in (b) would be produced, i.e. a flat area of exposed dentine (stippled) flush
with the surrounding enamel; such an appearance is seen on the antero-internal cusps
of the Piltdown molars. In (c) is shown a schematic section through a cusp partially
worn by natural attrition, illustrating the concavity of the dentine depressed below
the surrounding enamel. Artificial abrasion in the plane indicated would produce
the appearance shown in (d], i.e. a depressed " dimple " in the centre of a flat area
of dentine flush with the surrounding enamel ; such an appearance is seen on the
antero-external cusps of the Piltdown molars.

THE SOLUTION OF THE PILTDOWN PROBLEM 237

of dentine exposed to this degree form shallow concavities, because the less hard
dentine wears away more rapidly than the surrounding and harder enamel. The
possibility that the appearance of the occlusal surfaces of the Piltdown molar teeth
might be the result of an unusual type of natural attrition, which had proceeded so
rapidly that there was insufficient time for the exposed dentine to be hollowed out
below the level of the surrounding enamel, does not seem an acceptable explanation.
In the first place we have not been able to find, in an examination of 137 ape jaws
and 200 human jaws, a parallel condition in areas of dentine of similar extent exposed
by natural attrition. Secondly, the small punctate exposures of dentine on the
external cusps of the Piltdown molars are in fact already depressed below the surroun-
ding enamel, presumably as the result of natural wear (and thus indicating that the
latter was originally of the normal type) . Moreover, on the antero-external cusp of
both molars the punctate exposure of dentine is surrounded by a small and perfectly
flat area of dentine which, it appears, must have been exposed (like the areas of
dentine on the antero-internal cusps) by artificial abrasion (Text-fig. 3, c, d). These
flat areas of dentine are stained a brown colour and are sharply defined by a thin
outline of deeper staining. We have been able to produce outlines of somewhat
similar appearance in modern teeth which have been filed down and artificially
iron stained, and it seems probable that the outline of deeper staining may be due
to the relatively rich amount of organic substance at the dentino-enamel junction,
where there is a greater proportion of interprismatic substance to enamel-rods
(Noyes, 1948), and also to the considerable degree of branching and interlacing
of dentinal tubules at the surface of the dentine.

Another curious feature of the molars of the Piltdown mandible is that the dentine
on both Mi and M2 is much more extensively exposed on the antero-internal than
the antero-external cusps. But this is the reverse of natural wear, in which the outer
cusps of the lower molars are normally worn down more rapidly in the earlier stages
of attrition. Out of 200 human jaws there were only 4 in which (at an approximately
equivalent stage of wear) the dentine of a lower molar was exposed to an approxi-
mately equal extent on the outer and inner cusps ; in one other the area of exposed
dentine on the inner cusp was rather more extensive (though by no means to the
degree shown in the Piltdown teeth) . Thus with normal occlusion it must be at least
very exceptional to find human teeth in which the wear on the antero-internal cusps
is so much greater than on the antero-external cusps of Mi and M2 as to be com-
parable with the Piltdown molars. In ape molars also, so far as we have been able
to determine from the evidence of 137 jaws of the modern large anthropoid apes, as
well as 38 specimens representing 9 different genera of fossil apes, the wear is nor-
mally greater on the outer cusps. In no case did we find a lower molar tooth showing
the reversed condition seen in the Piltdown molars (though in one female gorilla
there were small punctate exposures of approximately equal extent on the antero-
internal and antero-external cusps of Mi). This evidence makes it difficult to
explain the relative wear of the outer and inner cusps of the Piltdown teeth except
by the hypothesis of artificial abrasion.

This inference is further supported by the sharp edges with no bevelling which
bound the flat occlusal surfaces of the molars at their margins. The absence of

238 THE SOLUTION OF THE PILTDOWN PROBLEM

bevelling at the external margin seems particularly significant, for normally this
margin shows distinct bevelling caused by the overlap in occlusion of the external
cusps of the upper molars. In only 10 out of 200 human jaws was the lateral margin
of the lower molar teeth found to be as sharp-cut as those of the Piltdown mandible
(but these exceptions did not show the other unusual features of the supposedly
fossil specimens). The sharpness of the edge in the Piltdown molars is consistent
with the postulate of artificial abrasion, and a similar appearance is to be seen on
the talonid basin of the second molar. This depression, which is still relatively
unworn, is also separated from the quite plane occlusal surface of the crown by a
sharp unbevelled edge. We have not been able to find a similar feature in our series
of ape and human jaws in which the lower molars have been exposed to natural
attrition. For, in normal occlusion, the protocone of the upper molar fits into the
talonid basin of the opposing lower molar and in normal wear produces a sloping
rounded edge at the margin of the depression. On the other hand, the sharp un-
bevelled edge seen in the Piltdown molar would be expected to result from an
attempt to produce a flattened surface by artificially planing down a more or less
unworn tooth. Examination of the occlusal surfaces of the two molars under a
binocular microscope provides further evidence, for here and there the enamel and
dentine are scored with extremely fine scratches, sometimes disposed in a criss-
cross pattern. Such scratches, which are not apparent to the same degree over the
enamel on the sides of the crowns, strongly suggest the application of an abrasive of
some sort.

A further point to notice in the first and second molars of the Piltdown jaw is
that they both show almost exactly the same degree of wear. But normally the
wear of the first molar is distinctly more marked, since it has been longer in use.
Exceptional cases, however, do occur in human lower molars in which the wear of
the two teeth is approximately equal, and it may happen that the second molar is
the more severely worn as the result of some defect in the upper dentition. In our
series of 137 ape jaws, we have only found three instances (chimpanzees) in which
the first and second molar teeth are worn to about the same degree. It appears,
therefore, that the condition in the Piltdown molars is at least very unusual.

One more suggestive feature is to be seen in the mandibular molars — their occlusal
planes are not quite congruous, i.e. they do not fit together to form a uniform
contour. The posterior margin of Mi at its inner end projects about 2 mm. above
the adjacent anterior margin of M2, and the occlusal plane of Mi is set at a slight
angle to that of M2. On the other hand, the outer end of the posterior margin of
Mi is level with the adjacent anterior margin of M2. The possibility that this might
be due to a post-mortem displacement of Mi, the occlusal surface of the latter having
as a result been rotated slightly outwards about the axis of its outer margin, is
negatived by two observations: (i) the radiograph of the molar teeth in lateral
view shows that their roots fit accurately into their sockets, and (2) there is no sign
of a contact facet on the exposed posterior surface of Mi (which might be expected
if this tooth had been displaced upwards relative to Mi). Radiographs of the teeth
in lateral view also appear to show the contact facets between them in accurate
apposition (see Text-fig. 2). It is true that the two molars are not in exact sym-

THE SOLUTION OF THE PILTDOWN PROBLEM 239

metrical alignment, but this is a common feature in an immature mandible in which
the eruption of the permanent dentition is in process of final completion and in
which the second molar has not completely rotated to its final position in the alveolar
border. On the other hand, a recent radiograph of the teeth from their occlusal
aspect shows a slight gap between the anterior root of Mi and the inner wall of the
socket, suggesting a slight displacement outwards of the tooth. Even so, it still would
not be possible with the slight readjustment which would be necessary to correct it,
to bring the occlusal planes of the two molars into precise conformity, the more so
because the posterior margin of Mi is slightly concave in contrast to the quite
straight anterior margin of M2. This evidence of the lack of conformity between
the occlusal plane of the two teeth adds further support to the hypothesis of artificial
abrasion, for it suggests that the process of paring down has been applied separately
to each tooth. Taken by itself, this is not perhaps entirely conclusive, for as the
result of mal-occlusion resulting from some defect of the upper dentition incon-
gruities may occasionally occur in teeth exposed to natural attrition.

The enamel bordering the dentine • exposure on the antero-internal cusps of Mi
and M2 has been cracked and chipped, and on M2 a small flake has evidently become
detached and been replaced in position with some adhesive. We have found in
our experimental grinding of molar teeth that this type of cracking and chipping of
the enamel is very liable to occur, though it can be minimized by embedding the
teeth in plaster of Paris, and it is therefore of particular interest to find that the
Piltdown molars are similarly affected.

The radiograph demonstrates the presence of a small lobulated odontoma in the
pulp cavity of Mi, an unusual feature which, however, has no particular relevance
to the Piltdown problem.

The isolated lower left molar (probably Mi) reported to have been found in a heap
of stones raked off a field two miles from the site of the Piltdown excavations is so
closely similar in dimensions and shape to the mandibular molars that it probably
belonged originally to the same individual. However, it does not show the same
degree of flat wear and, no doubt for this reason, some authorities have refused to
associate it with the Piltdown jaw. But if the wear of the molars has been artificially
produced, such an objection no longer remains valid. An examination of this isolated
tooth with a binocular microscope shows that the enamel on the occlusal surface of
the crown is scored with fine scratches, similar to, but rather coarser than, those
already noted on the mandibular molars.

The evidence of the isolated canine tooth found in 1913 is consonant with that of
the molars. The radiological evidence that this tooth had not yet completed its full
development appears to be sound. The pulp cavity is widely open at the apex of
the root, and even if this is assumed to be the result of post-mortem damage, it
would not account for the relatively large size of the cavity as a whole. But, if the
tooth is immature, it is difficult to explain the severe degree of attrition of the lingual
surface of the crown unless it has been artificially produced.1 For the entire thickness

1 The large size of the pulp cavity had been noticed many years ago by Lyne (1916), but he sought to
explain the combination of this feature with the severe wear on the assumption that the tooth is a
deciduous canine of unusual size.

24o THE SOLUTION OF THE PILTDOWN PROBLEM

of the enamel has been removed over the entire extent of the lingual surface, from
the anterior to the posterior border of the crown. Over a small area just above the
middle of the worn surface the dentinal wall of the pulp cavity has been reduced to
a thickness of less than i mm., and at one point the pulp cavity has actually been
penetrated. At that point a rather curious feature is seen in the radiograph, for it
appears that the opening into the pulp cavity has been plugged with some plastic
material which is not itself radio-opaque (but which contains some fine dust-like
particles which are radio-opaque).1 Further, the radiographs show no evidence of
the deposition of secondary dentine, with a narrowing of the pulp cavity, such as
might be expected if the severe attrition of its lingual surface had been naturally
produced. It is now clear that, in the original descriptions of the tooth, the material
used for plugging the opening into the pulp cavity was mistaken for secondary
dentine. Apart from the severity of the wear of the canine, the pattern of attrition
is quite unlike that found in any ape (recent or extinct) , whether the canine belongs
to the upper or lower dentition. It has, indeed, been argued that such a type of
wear might be theoretically possible in an ape's jaw, given certain unusual occlusal
relationships and movements of the jaw, but (apart from the questionable validity
of these arguments) the fact is that it has never been demonstrated to occur in any
known pongid or hominid. The contour of the worn surface is in fact peculiar, for
while it is evenly concave in a vertical direction it is almost flat from before back-
wards and it is not accompanied by an attrition facet on the anterior or posterior
margin of the tooth.2 It is exceedingly difficult to imagine an occlusal relationship
which could have produced such a contour by natural wear, and it is little wonder
that in the early discussions on the Piltdown " fossil " there was some controversy
whether the attrition was caused by contact with the opposing canine or lateral
incisor tooth. On the other hand, the contour of the worn surface is quite consistent
with the surmise that the latter was abraded by artificial means.

It may be argued that at least some of the details of the Piltdown teeth which
have been described, when considered separately, are inadequate by themselves to
confirm the thesis that the teeth have been deliberately fabricated to simulate fossil
specimens. But when they are taken together we are forced to the conclusion that
they could not possibly have been produced other than artificially.

In attempting to decide whether the jaw and teeth are those of an orang or a
chimpanzee, one must remember that the artificial abrasion has removed or other-
wise damaged the finer details of dental morphology which distinguish these two
genera. Even so, it is possible to say that they are almost certainly those of an orang.
Thus, the hypsodont character of the molars and the size and shape of their pulp
cavities are quite similar to those of orang teeth (Text-fig. 4), but differ markedly,
from those of chimpanzees which we have examined. Again, the pattern of the

1 The penetration of the pulp cavity was not evident in the radiographs of the canine reproduced in
the original communication of Dawson & Woodward (1914), for the reason that they did not happen
to have been taken in just the appropriate plane.

1 When the canine tooth was picked out of a heap of gravel in 1913, it was found accurately to resemble
in shape the plaster model of the canine which had already been reconstructed to fit the Piltdown
mandible. Commenting on the discovery in a postscript dated i6th September, 1913, Underwood
(1913) wrote " The tooth is absolutely as modelled at the British Museum."

THE SOLUTION OF THE PILTDOWN PROBLEM

241

dentine exposure corresponds very closely to that produced in our experimental
abrasion of orang molars, and the numerous small fissures and pits in the central
part of the occlusal surface are clearly the residual traces of crenulations of some
complexity. Indeed, in one orang jaw in which we abraded the first and second molar

TEXT-FIG. 4. Radiographs of (A) the Piltdown molar teeth, compared with (B) those
of the female orang mandible shown in PI. 27, fig. 2. The orang is evidently more
fully mature than the fossil specimen, as shown by the slightly smaller size of the
pulp cavities and narrower root canals of the molars, and by the fact that the mandible
as a whole is a little more robust. (For a radiograph of orang molars showing pulp
cavities of practically identical size and shape with those of the Piltdown teeth, see
Weidenreich, 1937, ng- 32°)- [X-rays by P. E. Purves (A) and G. M. Ardran (B).]

teeth, they were found to duplicate the appearance of the Piltdown molars to a
remarkable degree, not only in the extent and the contour patterns of the dentine
exposures on the several cusps, in the size, depth and abrupt margins of the central
basin, and in showing an approximately similar residuum of crenulations, but also
in the general proportions of the crowns as a whole (including their height as measured
above the enamel margin). See PL 27.

(II) THE " TURBINAL BONE "

In the supplementary note on the discoveries at Piltdown (Dawson & Woodward,
1914), Dawson stated " I saw two human nasal bones lying together with the remains
of a turbinated bone beneath them in situ. The turbinal, however, was in such bad

GEOL. II, 6.

20

242 THE SOLUTION OF THE PILTDOWN PROBLEM

condition that it fell apart on being touched, and had to be recovered in fragments
by the sieve ; but it has been pieced together satisfactorily". The only other
reference to this find was made by Woodward in the same communication, and is
limited to the following statement: " The remains of a turbinal found beneath the
nasal bones are too much crushed and too fragmentary for description ; but it may
be noted that the spongy bone is unusually thick, and has split longitudinally into
a series of long and narrow strips." No reason was given for identifying the fragments
as those of a turbinal (it is clear that a maxillo-turbinal bone is meant to be indicated
by this term), except presumably by implication its proximity " in situ " to the
nasal bones. Moreover, contrary to Dawson's statement, the tiny fragments (eight
in number) are separate and it is not possible to fit them together to form a complete
bony element.

From an examination of these fragments it is clear that, whatever their true
identification may be, they are certainly not those of a turbinal bone. They show
none of the characteristic features of the maxillo-turbinal (such as its extreme thin-
ness or its pitted and cellular texture). On the contrary, the fragments are relatively
thick and they show a longitudinally grained texture which indicates that they are
composed of Haversian systems arranged in parallel formation. Presumably, there-
fore, they are derived from the shaft of a limb bone (probably of some small animal),
but their precise identification is indeterminate.

3. THE PILTDOWN 'IMPLEMENTS"

By K. P. OAKLEY

Department of Geology, British Museum (Natural History)

(I) THE FLINT " IMPLEMENTS "

DAWSON recorded that the Piltdown gravel is composed of Wealden ironstone
pebbles, mixed to the extent of about one-sixth of the mass with angular brown
flints, a large proportion of which are tabular in form with fractured edges, frequently
conforming to the so-called eoliths of the Plateau Gravels. He was duly cautious
with regard to eoliths. " Whether natural or artificial, the fractures appear to
have been largely governed by the prismatic structure of the flint " (later he intro-
duced the term starch-fracture to describe breakage of this type). He pointed out
that the rolled examples had a deep iron-stained patina, whereas the unrolled ones
were less deeply stained and patinated. He said that there also occurred in the
Piltdown gravel " certain brilliantly coloured iron-red flints,, presumably more
highly oxidised than the prevailing flints, which were of a brown colour." He
continued : "Among the flints we found several undoubted flint-implements, their
workmanship being similar to that of the Chellean or pre-Chellean stage ; but in the
majority . . . the work appears chiefly on one face of the implements."

Only four of these " Palaeolithic implements " were figured by Woodward and
Dawson in their joint papers, as follows :

£.605. (PL 28, fig. 3B). Side-blow flake of pointed foliate form, 9 cm. X 5 cm.,
with maximum thickness 2*5 cm.

The whole surface is patinated and stained yellowish-brown, the edges are
dulled. See Dawson & Woodward 1913, pi. XVI, i, la, b; Sollas 1924, fig. 81 ;
Woodward 1948 : 40, fig. 90.

According to some unpublished notes, Woodward found this " best formed
tool " loose in one of the heaps of gravel rubbish at the main site.

£.606. (PI. 28, fig. 30). Triangular bifaced point, with area of smooth
brownish stained cortex at the butt ; 8 cm x 6 cm., with maximum thickness
of 2-5 cm. The flaked surfaces are patinated with a superficial pale yellowish
brown stain, the edges are dulled. See Dawson & Woodward, 1913, pi. XVI,
2, 2a ; Woodward 1948 : 40, fig. 8a.

This " palaeolithic tool " is recorded by Dawson (in Dawson & Woodward
1913 : 122, footnote i) as having been found in situ by Father Teilhard de
Chardin, in the loamy stratum of the Piltdown gravel (see p. 275).

£.607. (PI. 28, fig. 3A). Massive flake of squarish form, 14 cm. x 13 cm.,
with maximum thickness of 5 cm. The flaked surfaces are patinated and

244 THE SOLUTION OF THE PILTDOWN PROBLEM

stained bright orange, the edges are dulled. See Dawson & Woodward, 1913,
pi. XVI, 3, 3«, b ; Breuil, 1949, fig. 4. According to unpublished notes by
Woodward this specimen was found by Dawson on a heap of gravel rubbish
at the main site.

£.613. Heavily rolled, oval end-struck flake, 6-5 cm. x 5 cm., 1-5 cm. thick,
of dark greyish flint with thin buff patina, the surface of which has a patchy,
dark red ferruginous stain. See Dawson & Woodward, 1914 : 84, pi. XIV,
la-c ; Breuil, 1949 : 344, fig. i ; Oakley 1949, fig. 30.

This is the only artifact which was recorded as having been found in situ
in the dark brown basal gravel.

The first three specimens have a number of features in common :

(i) Although for the most part fractured by human agency, the form of each
appears rather more accidental than intentional ; (2) each is an atypical artifact,
not referable with certainty to any particular culture ; (3) each is in flint of a quality
not found as unworked pieces in the Piltdown gravel ; (4) the patina of each artifact
has a ferruginous staining which is unusual in its bright colour (varying from orange
to light yellowish brown) and in its patchy, rather stippled appearance (PL 28, fig. 4) ;
(5) each specimen shows evidence of some heavy localised battering (PL 28, fig. 3
A-c) on the original exterior or cortex of the flint prior to its being flaked from the
parent block.

The peculiar staining and the lithology of these three specimens are exactly
matched by a core which was discovered recently in a cabinet formerly belonging
to the late Mr. Harry Morris of Lewes. This specimen (PL 28, fig. 30) is now pre-
served in the British Museum (Natural History). In an accompanying note, Morris
drew attention to its similarity to the worked flints from Piltdown, and claimed
that to his own knowledge this flint had been artificially stained. The patina itself
appears to be natural, so the flaking is prehistoric, but judging from the style of work
it is probably Mesolithic or Neolithic rather than Palaeolithic.

When dilute hydrochloric acid is applied to the surfaces of the Piltdown " palaeo-
liths " and Morris's flint the orange or yellowish brown stain is dissolved and can
be wiped off, leaving a pale yellowish or greyish white surface of patina. In marked
contrast, it has been found that the brown patina of the " eoliths " and of other
flints collected from the Piltdown gravel, is unaffected by hydrochloric acid. (PL 28,
figs. 5, 6).

Experiments carried out in collaboration with Dr. A. A. Moss in the laboratory
of the Mineral Department of the British Museum have shown that the colour of
the orange and yellowish or reddish-brown stained flints from Piltdown can be
reproduced by dipping white-patinated flints in a solution of ferric chloride of
various strengths and then treating the wet stain in ammonia fumes to produce ferric
oxide.

The surface stains of a number of flints from Piltdown, including the figured ones,
were first tested for us by Dr. E. T. Hall in the Clarendon Laboratory, Oxford,
using his X-ray spectrographic method of analysis (Hall, 1953). The stains proved
to be entirely ferruginous, with the notable exception of the yellowish-brown stain

THE SOLUTION OF THE PILTDOWN PROBLEM 245

on E. 606 which contained in addition to iron appreciable traces of chromium (Table
I). This result was confirmed by Dr. A. A. Moss, using a direct chemical method of
analysis. Since chromium was not detected in the Piltdown gravel, or in any other
flints from the gravel, there can be no reasonable doubt that this implement has been
artificially stained.

TABLE I. — Composition of stains on Piltdown flints

Estimations by E. T. Hall
(in milligrams per sq. cm.)

Specimen Fe Cr

£.605 " Palaeolithic tool " . . . . 0-8 nil

E.6o6 " Palaeolithic tool (in situ) " . . . 0-5 0-2*

£.607 " Palaeolithic tool " .... 1-4 nil

£.613 " Palaeolithic flake " .... 1-6 „

£.685 Flint from surface . .... 1-2

£.956 "Eolith" 0-8

£.965 "Eolith" 0-5

£.985 "Eolith" 0-8

£.989 Palaeolithic flake-blade .... 0-2

£.2690 Morris flint ...... o-i ,,

Unreg. Sample of gravel . . . . . 0-9 „

* Chemical estimation gave circa o-i mg./sq. cm.

When a small chip was removed from the stained cortex of E. 606 for chemical
analysis, the staining was found to be superficial ; below its surface the cortex proved
to be pure white (PI. 28, fig. 30). Yet in brown flints normally found in the Piltdown
gravel the cortex is iron-stained throughout its thickness. Flint nodules with white
cortex do not occur naturally in the Piltdown district.

Apart from their orange or brownish staining the Piltdown " palaeoliths " could
quite easily be matched in the flint waste found at flint-mining or chipping sites
of Neolithic or later age on the Chalk Downs of Sussex. Dawson was perhaps not
far from the truth when he wrote of the Piltdown " palaeoliths " as follows : " They
resemble certain rude inplements occasionally found on the surface of the Chalk
Downs near Lewes, which are not iron stained " (Dawson & Woodward, 1913 :
122, footnote 2). The battered aspects of these specimens (PI. 28, fig. 3A-c), suggest
that they may have originated through fortuitous shattering of flint hammerstones
or anvils. Such accidentally fractured pieces are not uncommon on downland
flint workshop sites. Specimens E.6o5 and E.6o6 show no signs of use, which is in
accord with their being waste pieces. Some attempt has been made to shape two
of the edges of the thick flake E.6oy, but randomly broken pieces were sometimes
selected for use either as cores or as occasional tools by the Neolithic and Early
Bronze Age flint workers.

Thus it appears that the only flints found at Piltdown which Dawson and Wood-
ward considered worthy of figuring as probable palaeolithic implements show features
which are difficult to explain unless they were brought to the site from elsewhere and,
having been suitably stained, planted in the gravel for the excavators to find. The

246 THE SOLUTION OF THE PILTDOWN PROBLEM

fact that the pieces in question are crude and atypical (in the words of Ray Lan-
kester1 " unlike any known or denned industry ") suggests that they may have been
chosen so that they could be compared with the poorly denned and altogether dubious
artifacts which had then recently been found below the Red Crag of Suffolk (Moir,
1911). Whether in fact any artifacts occur in the Piltdown gravel is now doubtful.
No humanly fractured flint was noted during recent excavations (Toombs, 1952) ;
all the artifacts previously reported were either surface finds or were introduced
fraudulently.

A broken nodule of black flint with heavily bruised edges was found by Woodward
in the sandy layer overlying the basal gravel ; he suggested that it may have been
used as a hammerstone (Woodward 1917 : 2). In fact it bears more resemblance to
a broken paving cobble bruised by cart-wheels than to a prehistoric hammerstone.

(II) THE BONE " IMPLEMENT "

(PI. 29, figs. 7-10)

Next to the skull, mandible and canine tooth, the most remarkable find at Piltdown
(but not in situ) was an object shaped like the blade of a cricket bat, which had been
made out of a strip of bone from the femur of a large extinct elephant. Woodward
regarded it as possibly belonging to the Lower Pleistocene Elephas meridionals, but
it could equally well have come from one of the larger Middle Pleistocene species.
Dawson & Woodward (1915) regarded the object as an implement which had been
shaped by " a primitive tool, presumably a flint " when the bone " was in a compar-
atively fresh state." These conclusions were challenged at once when their paper
was read before the Geological Society. Reginald Smith said " the possibility of
the bone having been found and whittled in recent times must be considered," and
A. S. Kennard doubted whether the bone had been cut when it was fresh. No
comparable bone work has ever been found in known Palaeolithic industries and
recent experiments have shown (a) that the facets on the Piltdown specimen bear
no resemblance to the scratchy marks made on bone by a flint knife, nor, as Breuil
once suggested (1938), to the cuts of beavers' teeth, but must have been made by
an even-edged metal blade ; (6) that it is practically impossible to whittle a fresh
or recently dried bone, which can only be worked by flaking, scraping, sawing or
grinding ; and (c) that some fossilised bones, having more the texture of chalk, are
readily carved. Plate 29, fig. 10, illustrates a fossil bone from the Swanscombe
gravels which was whittled with a steel razor and then stained with a ferric solution
and varnished to reproduce as nearly as possible the present appearance of the cut
surfaces of the Piltdown specimen.

In conclusion, the Piltdown bone " implement " is a piece of the femur of a fossil
elephant, obtained probably in two weathered pieces from a Middle Pleistocene
brickearth or sandy formation. The ends were whittled with a steel knife, and the
newly cut surfaces were stained with an iron solution. The small fragments of
bone, now chemically shown to be from the same original source, which were found
in the basal clay suggested to the excavators that the implement belonged to that
level (see p. 253).

1 In lit. to A. S. Woodward, 3ist Jan., 1913.

4. THE PILTDOWN MAMMALIA

By K. P. OAKLEY

Department of Geology, British Museum (Natural History)

BETWEEN 1911 and 1914 eighteen fossil mammalian bones and teeth were found at,
or in the immediate vicinity of, the Piltdown skull site. Four were recorded as
having been found in situ in or below the chocolate-coloured basal gravel, two on
the surface of the adjoining field and the remainder on the spoil heaps at the edge
of the small pit. Considering the thinness of the gravel (average thickness 18 in.),
the small size of the pit (less than 50 x 10 yards in area), and the extreme rarity
of fossils in the Pleistocene river gravels of Sussex, this was a remarkable yield.

The fossils are heterogeneous in character, ranging from a heavily rolled enamel
cap of a Mastodon tooth to a large piece of cervid antler with almost undamaged
surface. Apparently they were not scattered through the gravel but were confined
to one or possibly two pockets. Woodward (1948) wrote :

" After Mr. Dawson's death in 1916, I was able to open a series of pits along the
other side of the hedge in a field adjacent to the original pit. There I was helped by
Professor Elliot Smith . . . and others. We began close to the spot where the skull
was found and worked in both directions from that place. Our efforts however
were all in vain. We found nothing of interest in the gravel."

In 1950 a further trench, 4 ft. x 32 ft., was dug in undisturbed gravel by the
Nature Conservancy, and all the excavated material was sieved and carefully
examined, but no bones nor teeth were found (Toombs, 1952).

Some of the Piltdown fossils (" Stegodon", Mastodon arvernensis and Rhinoceros
cf. etruscus) are undoubtedly Villafranchian (" Upper Pliocene " of earlier authors,
but now classed as Lower Pleistocene) ; but others are not older than Middle or
Upper Pleistocene. When they were first described there was some question as to
whether gravels of two ages were present. Dawson concluded : " It is clear that
this stratified gravel at Piltdown is of Pleistocene age, but that it contains in its
lower stratum, animal remains derived from some destroyed Pliocene deposit
probably situated not far away, and consisting of worn and broken fragments"
(Dawson & Woodward, 1913 : 123). In a later paper he wrote : " We cannot resist
the conclusion that the third or dark bed is in the main composed of Pliocene
drift ..." (Dawson & Woodward, 1914 : 85).

(i) " Derived " or Villafranchian Group

With the increase of knowledge, the presence of " Pliocene " (Villafranchian)
mammalian remains at Piltdown became increasingly difficult to explain, for, with
one significant exception (see Table IE footnote), none has been found elsewhere in
southern England. If they were to be found in situ in Sussex, they would be expected
in terrace or plateau deposits not less than 200 ft. above the level of the Ouse, but
Edmunds (1926 : 68) demonstrated that the Piltdown gravel is part of a terrace
50 ft. above the Ouse (see p. 273). Thus it seemed that they could only have come

248 THE SOLUTION OF THE PILTDOWN PROBLEM

from a block of indurated fossiliferous sand which worked its way down from a
higher level and disintegrated on the Ouse flood-plain in 5o-ft. terrace times.

When it had been established that the Piltdown mandible was a forgery it became
probable that the Villafranchian fossils were also introduced in order to suggest that
" Piltdown Man " dated from Pliocene times.

In his original description of one (£.596) of the three pieces of molar tooth referred
to " Stegodon " Woodward wrote : " It cannot be referred to the upper Pliocene
Elephas meridionalis , because in this species the valleys are deeper in proportion to
their width, while the ridges are more plate-like and parallel in their upper portion.
The new specimen is, therefore, of an earlier Pliocene type which is best known from
the Siwalik Formation in India and has not hitherto been found in Western Europe "
(Dawson & Woodward, 1913 : 142). Later, the Piltdown specimens were compared
to Elephas planiforns, of the Upper Siwaliks, now regarded as Lower Villafranchian.
There are no specimens comparable with E. planifrons from the English Red Crag
in the British Museum nor in the Geological Survey collections. That three pieces
of Elephas cf . planifrons should be recorded at Piltdown might therefore be regarded
as very remarkable. In their reddish colour the Piltdown pieces resemble Red Crag
fossils, and also closely match the Piltdown hominoid remains. This similarity in
colour led some investigators to conclude that in spite of their unrolled condition
the Piltdown skull and mandible were of Villafranchian age (Hopwood, 1935).

If the " planifrons " molar fragments were introduced at Piltdown it is probable
that they were obtained from some foreign source and artificially stained to match
the Piltdown cranial bones and mandible. The Piltdown mandible was given a rich
mahogany colour by a process which involved the use of iron and chromium com-
pounds, and it appeared likely that the same method would have been applied to
imported "planifrons" teeth. Samples of the iron-stained cementum of the two
critical specimens from Piltdown (£.596 and £.620) were analysed spectrographi-
cally in the Department of the Government Chemist by Mr. H. L. Bolton, who
reported that they contain significant traces of chromium (circa 0-3 and 0-1%
respectively).

Any remaining doubt that these pieces of "planifrons" molars were of foreign origin
was dispelled by their radioactivity. In the hope of tracing the origin of these pieces
a series of mammalian teeth from the main Villafranchian localities was included
among the fossils selected for the tests (p. 276). The results obtained reinforce the
conclusion that the Piltdown specimens were not obtained from an English deposit.
The uranium content of fossil teeth, while increasing with geological age, is subject
(as fluorine is) to considerable fluctuation from place to place. Nevertheless it
appears that within relatively uniform strata of limited extent the ratio of the
extremes of the variation in comparable material does not generally exceed the value
of three. Thus a specimen consisting largely of dentine or cementum with a radio-
activity of 200 net counts per minute is extremely unlikely to have come from a
deposit such as the Red Crag in which random samples of teeth show a maximum
radioactivity of less than 30 c.p.m. Uranium (again like fluorine) is adsorbed more
readily by cementum and exposed dentine than by enamel, and on an average, the
radioactivity of enamel in a fossil tooth is only about one-third that of dentine or

THE SOLUTION OF THE PILTDOWN PROBLEM 249

cementum (PI. 30, fig. 13). The radioactivity figures in Table II are maxima based
on samples which were mainly cementum and dentine.

TABLE II. — Radioactivity of Piltdown elephant teeth compared with mammal teeth from

Villafranchian sites

c.p.m.
Source of specimen (Max.)

" Piltdown " (£.620) 355

" Piltdown " (£.596) 203

" Piltdown " (E.597) 175

*" Portslade " (M.I0436) 28

Red Crag, Suffolk 15

Red Crag, Suffolk 13

Red Crag, Suffolk 12

Red Crag, Suffolk 12

Red Crag, Suffolk 9

Red Crag, Suffolk 6

Doveholes, Derbyshire ...... 25

Netherlands (dredged specimen, ? leached) . . . i

Seneze ....... 15

Puy-de-Dome . . . . . . • . <io

Chagny-Bellecroix ....... 5

Val d'Arno 26

Siwaliks ....... 19

Siwaliks ......... 8

Algeria ......... 18

Algeria ......... 15

Morocco . . . . . . . . . <io

Ichkeul, Tunisia . . . . . . 195

* About 191 1 the late Dr. Eliot Curwen obtained some fossils from workmen digging brickearth at
Portslade, near Brighton, Sussex, including a well-preserved, grey to buff -coloured, molar of Ursus
arvernensis (a Villafranchian species of bear) and two complete pale buff-coloured limb-bones of an
undetermined species of the same genus, which he presented to the British Museum (Nat. Hist.) where
they are registered under M. 10436 and M. 10571-2. See White 1926, pp. 85-6. The radioactivity of
these three specimens proved to be higher than any of the British Pleistocene fossils which have been
tested (the Piltdown specimens exluded). Their Lower Pleistocene age therefore seems assured, and
since they are not in the condition of derived specimens they can have no natural place in the low-
level Portslade brickearth which is Upper Pleistocene. In striking contrast a mammoth molar from the
same site and undoubtedly contemporary with the brickearth proved to have a very low radioactivity.
There is a strong suggestion that the specimens of Ursus had either been " planted " or had been traded
by workmen who happened to have access to some collection of foreign fossils.

Fossils from nearly a dozen Villafranchian localities in Europe and Asia were
tested, but none showed radioactivity in excess of 28 c.p.m. Professor C. Arambourg
enabled us to extend the range of comparison by generously providing specimens
from North African localities. A molar tooth originally recorded as Elephas cf.
planifrons, from Garaet Ichkeul in Tunisia, has proved to have a radioactivity closely
comparable with the Piltdown specimens. Not only is the count rate in the Ichkeul
specimen and £.596 from Piltdown almost identical, but the difference between the
activity of the enamel and of the cementum is unusually small in both. There is
also close agreement in the fluorine content of these specimens. Ichkeul, 20 km.
south west of Bizerta, is a richly fossiliferous locality, at which Elephas cf. planifrons
is stated to be the commonest species represented (Arambourg & Arnould, 1950 : 155).

250 THE SOLUTION OF THE PILTDOWN PROBLEM

%F p.p.m.

— v Tnr» r r

°/ P O />TT O

/0-t2^6 e*~>3V$

_., , ,. /"dentine and cementum 8-2 . 610

Piltdown' plamfrons molar £.596^^ .... I>8 .520

., f dentine and cementum . 7-9 . 1060

Piltdown plamfrons molar E.62O<

\^ enamel . . . . 2-6 . 170

f dentine and cementum . . . 8-4 . 580

Tchkeul plamfrons molar <

\enamel . . . . . 2-7 . <48o

f dentine and cementum . . . 7-1 > 56

Siwahk plamfrons molar-; .

\^enamel ...... 2-0 . 45

Siwalik plamfrons molar, dentine and cementum . . . 7-3 >24

Proof that three Villafranchian mammalian teeth were introduced into the Pilt-
down gravel suggests that the others were also introduced. The molars of Mastodon
and Rhinoceros showed no evidence of artificial staining, but to judge from their
colour, mineralization and radioactivity, it appears probable that they were ob-
tained originally from the Red Crag of East Anglia.

(ii) " Contemporary " Group

Division of the Piltdown mammalia into a derived Villafranchian group and a later
group contemporary with the deposition or re-arrangement of the gravel was mainly
based on the known ranges of the genera and species represented ; colour and state
of mineralization of the specimens were also taken into consideration. None of these
criteria proved altogether satisfactory. Hopwood (1935) divided the specimens into
a dark-coloured, heavily mineralized group, comprising the teeth of " Stegodon "
(Elephas cf. planifrons), Mastodon, Rhinoceros and Hippopotamus, and a paler, less
mineralized group, including the remains of Equus, Cervus and Castor. He pointed
out, however, that Hippopotamus had never been recorded in England from deposits
earlier than the Cromer Forest Bed and that the broken edges of the molar from
Piltdown were sharp, suggesting that it was not a derived specimen. Moreover,
the edges of two of the fragments of " planifrons " molars were also sharp, and since
there could be no question of the geological age of these pieces, it had to be admitted
that some Villafranchian specimens had found their way into the Piltdown gravel
without being rolled. This important observation suggested that the dark-coloured
but unrolled pieces of the Piltdown skull could belong to the older or Villafranchian
fauna. This had been the opinion of E. T. Newton, and was admitted as a possibility
by Dawson (in Dawson & Woodward, 1913 : 151), although he was more inclined to
regard " Eoanthropus " as contemporary with the later, Pleistocene fauna.

The fluorine-dating method seemed a suitable means of checking the relative ages
of the Piltdown remains, and the results (Oakley & Hoskins, 1950) did agree broadly
with the palaeontological dating of the mammalian remains. All the undoubted
Villafranchian teeth (Elephas cf. planifrons, Mastodon, and Rhinoceros cf. etruscus]

THE SOLUTION OF THE PILTDOWN PROBLEM 251

showed a very high fluorine content (i •9-3-1%), whereas the remainder, all of which
could be post-Villafranchian, showed considerably less (o-i to 1*5%). But the range
of fluorine content in the probably post-Villafranchian specimens was far greater
than was to be expected in fossils of a single age-group, and it was therefore inferred
that the Piltdown gravel had been re-arranged, and new mammalian remains
introduced, on several occasions.

According to the results obtained, all the " Eoanthropus " material contained on
an average the same small amount of fluorine as the bones and teeth of Castor. This
seemed to confirm Dawson's opinion that the beaver remains were the only fossils
from the pit that were contemporary with " Piltdown Man " (Dawson & Woodward,
1914 : 86).

One other fossil recorded from the Piltdown gravel showed an equally low fluorine
content : the molar tooth of Hippopotamus, which Hopwood placed in the Villa-
franchian group on account of its dark colour. Its enamel is colourless and almost
unaltered, but its dentine, containing < 0-05% fluorine, is stained brownish black
throughout. The low fluorine content appeared to indicate that it was not derived
from an older geological formation, but belonged to the latest faunal group in the
gravel. Yet its preservation is quite unlike that of the associated teeth of Castor.
Considering that the " planifrons " molars from Piltdown had been artificially
stained and then introduced at the site, the colour of the Hippopotamus molar
suggested that it, too, had a similar history. This was confirmed by spectrographic
analysis carried out in the Department of the Government Chemist by Mr. H. J.
Dothie, who found that the dark-coloured dentine contained i% of chromium, a
clear indication of artificial staining.

Drs. Weiler and Strauss (see below, also Table V) showed that this tooth has
an unusually high ash content, and a correspondingly low organic content. This and
its exceptionally low fluorine content suggests that it could only have come from a
limestone cave deposit, in which fluorination of bones and teeth is usually minimal,
Hippopotamus has only somewhat rarely been reported from British caves, but
occurs abundantly in the calcareous cave deposits of the Mediterranean islands.
Some of the molars from the Ghar Dalam Cave in Malta correspond in size with the
Piltdown specimen, and have a low fluorine content combined with a low organic
content. The Maltese teeth are creamy white in colour. Experiments in the Depart-
ment of Minerals showed that by soaking one of the Maltese specimens in a solution
of ferrous sulphate or iron alum, precipitating the iron as ferric hydroxide, and
then treating with tannic acid to produce a blackish tinge the colour of the
Piltdown molar could be reproduced exactly.

Percentages

N C H20 f/P*°5 CaCo3 Ash P*^

(X 100) 0U3 O

Dentine of Recent teeth . >2 >6 >i3 <o-3 5 <6o <i

Piltdown hippo molar dentine 0-06 2 12 0-3 2 84 3

Ghar Dalam hippo molar ,, nil nil 7 0-3 14* 87 7
* After staining in an iron sulphate solution this was reduced to 5 per cent.

252 THE SOLUTION OF THE PILTDOWN PROBLEM

The presence of chromium in the specimens from Piltdown suggests that they were
treated with chromic acid or a dichromate solution with the idea of aiding the
oxidation of a ferrous staining agent. If an iron sulphate had been used, it might
account for one other feature of the composition of the Piltdown Hippopotamus
molar, namely, the presence of calcium sulphate.

In 1949 an X-ray powder diffraction photograph of the blackened dentine, taken
in the Government Chemist's Department, revealed the presence of calcium sulphate.
Using the same technique Dr. G. F. Claringbull later detected calcium sulphate
(gypsum), apparently partly replacing the calcium phosphate, in several of the
bones and teeth from Piltdown. With Dr. M. H. Hey he has shown (p. 268) that
the gypsum could be the result of artificial treatment of the specimens, and probably
originated through interaction of the calcium phosphate of the bone or dentine and
a solution of an acidic iron sulphate.

Tests carried out by Dr. C. Bloomfield of Rothamsted Experimental Station
showed that, at the present day, SO3 (sulphate) ions are unusually low in the Piltdown
gravel, hence it is not possible to maintain that the sulphate-bearing specimens owe
their composition to some natural mineralizing process peculiar to the deposit.
No gypsum crystals could be detected by X-ray analysis of the fine fraction of the
gravel.

Eventually samples of all the Piltdown mammalian specimens were examined for
the presence of chromium, either spectrographically in the Department of the
Government Chemist, or chemically in the Department of Minerals of the British
Museum, where they were also submitted to X-ray crystallographic analysis (see
p. 269).

Several of the results obtained are worth commenting on here. Chromium and
gypsum were detected in a premolar of Hippopotamus (the first Piltdown fossil
shown to Woodward), proving that it too had been artificially stained. This tooth,
having a higher fluorine content (1-1%), presumably did not come from the same
source as the molar. But both the hippopotamus teeth are less radioactive than any
Pleistocene fossils from sand or gravel that have been tested (see Table XI), sug-
gesting that they are from calcareous or argillaceous deposits, for it is known that
phosphates are less prone to adsorb uranium where the strata are rich in calcium
carbonate or consist largely of clay (Davidson & Atkin, 1953 : 27).

Chromium was also found in the longitudinally split portion of a Cervus metatarsal
(Dawson & Woodward, 1913 : 121, 142), which had been included in the so-called
contemporary group. No sulphate was detected in this bone, indicating that the
staining solution used either was not sulphate, or was not sufficiently reactive to
cause replacement of the phosphate. A few specimens in the " contemporary "
group, including the mandible and incisor of Castor fiber, were found to contain
calcium sulphate, but no chromium. These were presumably stained by another
technique, which dispensed with the use of a dichromate solution as oxidizer.

The fluorine and organic content of the beaver mandible could be matched among
bones from either Late Pleistocene or Holocene deposits.

In addition to the artificially stained mandibular fragment and incisor, the
Piltdown collection includes two molars of Castor fiber. They both show superficial

THE SOLUTION OF THE PILTDOWN PROBLEM 253

iron-staining which may be artificial. One of the molars was left in its gravel matrix.
This seemed to be visible proof that the Piltdown gravel did in fact contain fossils,
but on examination the block containing the tooth proved to be an artifact. When
soaked in water it began to break down, revealing that it consisted largely of loamy
material with a concentration of pebbles on the outside. When the water in which
the block had been soaking was evaporated there was a large amount of gummy
residue (0-8 gm.). According to its former label one of the pieces of " Stegodon "
teeth (£.620) was also kept for a time in a similar lump of hardened gravel (see also
Dawson & Woodward, 1914 : 84).

One of the fragments of a cream-coloured bone said to have been found in situ
in clay at the base of the Piltdown gravel (and regarded by Dawson and Woodward
as indicating the source of the worked slab of elephant femur) is still embedded in
a lump of loam, adhering to the middle of a slab of ironstone. On close examination
the loamy matrix shows every indication of being faked. It contains small scattered
pebbles set at various angles, and its shows cracks and bells of burst air-bubbles
such as appear if loamy matter is worked into a paste and then allowed to set.
The sliver of bone embedded in this artificial matrix is identical in composition with
the worked elephant bone which, to judge from its state of mineralization and radio-
activity, could have been obtained from one of the brickearths in the 5o-ft. terrace
of the Thames or from an equivalent deposit in the Somme Valley.

% F p.p.m.

x 100

%P205 eU308

Elephas femur (worked), Piltdown . . . 4-2 . 10

Fragment from " base of gravel," Piltdown . 4-2 . 9

Cervus antler, Piltdown . . . . . 5-4 . 1 1

Elephas metatarsal, Crayford Brickearth . . 8

Rhinoceros ulna, Crayford Brickearth . . 3-5

Elephas astralagus, Ilford Brickearth ... 33

Rhinoceros astralagus, Menchecourt Sands, Somme 3-4 . (to)*
* Estimation based on an associated specimen.

The base of an antler of red deer, Cervus elephus, and an upper molar of Equus cf.
caballus (Irving, 1913) were found together in the adjacent field to the west of the
gravel pit, " on the surface close to the hedge " (Dawson & Woodward, 1913 : 121).
Neither bears any sign of having been rolled on the river bed. In its fluorine content
and radioactivity the stag antler is closely comparable with the worked elephant
bone. The horse tooth has a much lower fluorine content and shows no radio-
activity, indicating that it was derived from a different deposit. The surfaces of
both the horse tooth and the stag antler show patchy red iron-staining which recalls
in appearance the artificially stained flints (p. 244).

Of the eighteen specimens of fossil mammals recorded from the Piltdown gravel
by Dawson and Woodward, ten are unquestionably frauds, and there are strong
grounds for believing that this is also true of the remainder. Since the gravel is
decalcified (pH 6.5) it is probably unfossiliferous.

5. THE COMPOSITION OF THE PILTDOWN

HOMINOID REMAINS

•

By K. P. OAKLEY

Department of Geology, British Museum (Natural History)

(i) THE MANDIBLE AND TEETH

THE discovery in the autumn of 1953 that the Piltdown mandible contained the
same amount of nitrogen as fresh bone might be regarded as proving its modernity
beyond all doubt. In actual fact, however, this would not have been conclusive
evidence without the cross-check provided by fluorine analysis. Thus, an ulna of
woolly rhinoceros (M. 12575) found at a depth of 42 ft. in an Upper Pleistocene river
deposit on the site of Lloyd's in the City of London (Warren Dawson, 1925) has a
nitrogen content about the same as that of the Piltdown mandible. The reason for
the remarkable preservation of the rhinoceros bone is that it was embedded in an
unoxidized clay — an environment in which bone-protein decays very much more
slowly than in the oxidizing environment of sand or gravel. In marked contrast, a
fragment of mammoth femur (M. 12946) found at a depth of 20 ft. on the same site,
and of the same general age, but preserved in sand, was found to contain very little
nitrogen. Fortunately the fluorine content of bone increases at about the same rate
in sand (or gravel) and in clay. Thus the fluorine content of the rhinoceros bone is
almost the same as that of the mammoth bone.

% N % F

t TI j< -i {Rhinoceros, in clay . av. 3-4 . i-i
Upper Pleistocene bones from Lloyd s site < „, . , . , J

\^Elephas, in sand . o • i 1-3

Piltdown mandible, " in gravel " . . . . . . 3 '9 • <o-O3

Fresh mammalian bone . . . . . . . . 4'1 • °'°3

If the Piltdown mandible had occurred naturally in the shallow Piltdown gravel
(early Upper Pleistocene) it should not have contained more nitrogen than the
skull bones. Moreover, its fluorine content should have been greater than that of
fresh bone no matter whether the matrix were clay or gravel.

In one account of finding the mandible Woodward (1948 : n) wrote : " It had
evidently been missed by the workmen because the little patch of gravel in which it
occurred was covered with water at the time of year when they reached it." At an
early stage in the present investigation it had to be borne in mind that conditions
in the Piltdown gravel might have been exceptional and that reducing conditions
in the basal bed had preserved collagen in the bone. Through the courtesy of
the Director of the Rothamsted Experimental Station the chemical conditions in

THE SOLUTION OF THE PILTDOWN PROBLEM 255

the Piltdown deposits were recently examined by Dr. C. Bloomfield, who found that
the "redox potential" in the basal bed indicated oxidizing, not reducing, conditions.

In March, 1952, Professor J. T. Randall and Dr. A. V. W. Martin undertook to
examine the collagen in samples of the teeth by means of the electron-microscope.
The first results were inconclusive, for in drilling the samples frictional heat had
probably de-natured the collagen. Early this year a new attempt was made to
determine the state of collagen in the mandible. A small piece of the bone was sawn
out and submitted to Dr. Martin, together with a similar sample of the cranium
and other selected controls. Electron-micrographs of the decalcified residue of the
mandible sample revealed the presence of fairly well preserved collagen fibres with
characteristic banding at intervals of 640 Angstrom units (PL 30, fig. n), whereas
electron-micrographs of samples of the skull showed no trace of collagen fibres. A
residue of the Lloyd's rhinoceros bone also proved to contain collagen fibres, but
they were partly de-natured and showed only vague shadows of the original
banding. The only fossils in which collagen fibres have been found previously are
from frozen ground — see Randall et al., 1952. Collagen is denatured at 70°-ioo° C.

It has been suggested that collagen fibres may be preserved intact through the
action of peat acids, but none was detected in a sample of human bone with un-
diminished nitrogen content from Holocene peat at Branston, Nottinghamshire, nor
in a sample of human skin from the Tollund Bog, Denmark (kindly supplied by
Dr. Knud Thorvildsen of the National Museum Copenhagen).

Estimation of the fat content of the mandible, suggested by Heizer & Cook (1954 :
94) as a possible means of confirming its modernity, was considered impracticable
with a sample of the limited size available for such a test.

In addition to the nitrogen content, the organic carbon, water and ash contents
of the mandible have been determined and compared with those of selected controls.
The results confirm that this bone is modern, and also show that such artificial
treatment as it received has affected the organic fraction only very slightly.

%N %C %H20 %Ash

Modern ungulate limb-bone . . . (4-0) . 14-0 . 24-5 . (53)

„ „ „ 2nd sample. (4'i) • IO'3 . 21-2 . — •

Piltdown mandible . . . . (3-9) . 14-5 . 25-0 . (61)

Piltdown (left parietal) . . . . 1-4 . 6-1 . 17-8 . 62

Neolithic human skull, Coldrum . . 1-9 . 6-3 . 18-2 . 71

Lloyd's rhinoceros ulna . . . 3-4 . 10-4 . 18-9 . (67)

Lloyd's mammoth femur . . . O'i . 2-6. 10 -8. (8~)

Note. — The figures in brackets are determinations made on separate samples and therefore inde-
pendent of other figures on the same line.

The dark mahogany colour of the mandible, matching that of the skull bones
very closely, is relatively superficial. Drilling revealed that the interior tissue is
buff to pale grey in colour, suggesting that the organic matter filling the pores of
the bone prevented the iron-staining solution from penetrating deeply. When the
mandible was being drilled with a dental burr to procure an adequate sample for
the re-determination of fluorine, there was an odour of burning, and the ejection

256 THE SOLUTION OF THE PILTDOWN PROBLEM

consisted of minute shavings. When the skull bones were drilled in the same
way there was no odour, and the sample consisted of powder.

The mandible shows practically no radioactivity (see Tables X, XII), which is a
further confirmation of its modernity.

The fluorine content of the canine and of the molars in the mandibular ramus was
estimated in 1949 as < 0-1%, but as the mandibular bone itself appeared to contain
c. 0'2%, and as the probable experimental error on samples of the small size then
tested was known to be about ± 0-2% (Oakley & Hoskins, 1950, 379-80), there
seemed no reason to regard the canine or the mandible as more recent than the
human cranium, with fluorine content estimated to vary from o-i to 0-4%. In 1953,
new samples of the teeth and of the skull bones were submitted to the Department
of the Government Chemist, where Mr. C. F. M. Fryd had devised a technique for
estimating smaller amounts of fluorine than could be measured in 1949. The experi-
mental error in the determination of fluorine obviously depends on the size of the
sample and the amount of fluorine it contains. The fluorine content of the Piltdown
skull bones was determined in 1953 as 0*14 to 0-18%, and the limits of error as
± 0-02%. Where the amount measured was exceptionally small, the fluorine
content was recorded as less than o*ox%, the true figure lying between O'Ox and zero.
In 1953 all the determinations of the fluorine content of the Piltdown mandible and
teeth proved to lie between 0-04% and zero. These results indicated that whereas
the skull bones were probably prehistoric, the canine tooth, the mandible and the
isolated molar were modern. This conclusion was reinforced by comparing the
nitrogen content of the Piltdown bones and teeth (dentine) with that of modern
and fossil specimens.

In order to eliminate any possibility that the nitrogen found in the dentine
samples was not original but due to contamination of the samples, their organic
carbon was also determined. The carbon /nitrogen ratio in the molars and in the
canine proved to be slightly lower than in the dentine of two modern teeth which
were used for comparison, but approximately the same as in the organic matter of
bone (2-296 i 0-266, Cook & Heizer, 1952 : 4).

% N % C C/N

Modern beaver molar (old individual) 2-2 . 6-5 . 3-0

Modern orang-utan canine . . 3*9 . 12-8 . 3-2

Piltdown canine . . . . 5-1 . 12-1 . 2-3

Molars in Piltdown mandible . 4-3 . 10-0 . 2-3

Isolated Piltdown molar . . 4-2 . 10-7 . 2-5

In 1950 Dr. David B. Scott of the National Institute of Dental Research, Bethesda
(Maryland) undertook to examine collodion replicas of the surfaces of the Piltdown
teeth, using the metal-shadowing technique which he has developed with Wyckoff
(1946). After examining replicas of the buccal surfaces of the molars in the Piltdown
mandible, Dr. Scott reported that " they are not readily recognizable as ancient
teeth, since they show very little evidence of post-mortem damage". But, in contrast,
replicas of the isolated molar and of the crown of the canine near the tip revealed

THE SOLUTION OF THE PILTDOWN PROBLEM 257

considerable damage. These findings agree with the results of the present detailed
re-examination, that the molars in the mandible have been artificially abraded only
on the occlusal surfaces, whereas in the canine and isolated molar the buccal surfaces
also have been smoothed artificially.

The black coating on the canine is a paint made from a natural bituminous earth
containing iron oxide, probably Vandyke brown (see p. 272). If bituminous matter
were not out of place in a highly oxidized gravel it might have been regarded as a
natural incrustation. It should also be recorded that Dr. Claringbull found a minute
spherule of an iron alloy embedded in the coating on the labial surface of the crown.

The pulp cavity of the canine contains 19 loose sand grains, mostly radio-opaque.
Some were extracted for examination and proved to be pellets of limonitic iron-
stone identical with those which occur in the sand-fraction of the Piltdown gravel.
As seen in the radiograph of this tooth (Weiner, Oakley & Le Gros Clark, 1953, pi. 9,
fig. 4), all the grains are 1-2 mm. in diameter. If they had been naturally washed
into the cavity finer grade material would be expected to have entered with them,
for 30% of the grains in the sand-fraction of the Piltdown gravel are less than I mm.
in diameter. The cavity has been sealed by an ovoid grain of hard limonitic iron-
stone tightly wedged into the aperture of the truncated apex.

(II) THE PILTDOWN SKULL BONES

As the fluorine and nitrogen content of the cranial bones were consistent with their
being fairly ancient, it seemed at first that the hoax had been based on a genuine
discovery of portions of a skull in the gravel, and that the animal remains and
implements had been subsequently " planted " to suggest that it was Pliocene or
Early Pleistocene in age. As the investigations proceeded the skull too became
suspect. Dr. G. F. Claringbull carried out an X-ray crystallographic analsyis of
these bones and found that their main mineral constituent, hydroxy-apatite, had
been partly replaced by gypsum. Studies of the chemical conditions in the Piltdown
sub-soil and ground-water showed that such an unusual alteration could not have
taken place naturally in the Piltdown gravel. Dr. M. H. Hey then demonstrated
that when sub-fossil bones are artificially iron-stained by soaking them in strong
iron sulphate solutions this alteration does occur. Thus it is now clear that the
cranial banes had been artificially stained to match the gravel, and " planted " at
the site with all the other finds. The presence of chromium in some of the bones is
now more readily explicable, for a dichromate solution might have served to aid
the oxidation of iron salts used in staining the bones.

Since all the " local Upper Pleistocene " fossils with comparable composition
have proved to be fraudulent introductions, the low fluorine content of the skull
indicates that it is more probably post-Pleistocene than Pleistocene in age.

In 1912, no organic matter could be detected in the small piece of the Piltdown I
calvaria submitted for analysis to Mr. S. A. Woodhead, Public Analyst of East
Sussex (Dawson & Woodward, 1913 : 121). The specific gravity of the powdered
fragment was also measured (2 -115) ; but neither of these determinations was
significant so long as no comparison was made with the mandible. The first physical

GEOL. II, 6. 21

258 THE SOLUTION OF THE PILTDOWN PROBLEM

comparison between the mandible and the calvaria fragments was recorded in a
note dated 1925 by Dr. A. T. Hopwood, who found that their specific gravities,
measured in vacuo, were as follows :

Mandible 2-06 ; Piltdown I occipital 2-13 ; Piltdown II frontal 2-18.

The specific gravities of the Piltdown II occipital and isolated molar have been
determined more recently as 2-20 and 2*18 respectively. If allowance is made for
the density of the molars, the specific gravity of the bone of the mandible becomes
2-05. The difference between the specific gravity of the mandible and of the calvaria
would possibly have been greater if the constituent apatite had not been so extensively
replaced by gypsum, which is a lighter mineral, although additional iron oxide may
have counterbalanced this effect.

The age of the Piltdown skull has been questioned on the score that it included
nasal bones in close association (Marston 1950 : 293). Unless ankylosed before death
the nasal bones have commonly separated even in recent burials. It was therefore
difficult to understand their occurrence together in the upper disturbed layer of the
gravel (Dawson & Woodward, 1914 : 84). However, there was always the possibility
that the nasal bones did not belong to the Piltdown skull. These bones show partial
replacement by gypsum, indicating that they were artificially iron-stained. To judge
from their composition they were not obtained from the same source as the other
cranial fragments. In drilling samples from the nasal bones the ejection consists of
shavings (as when the Piltdown mandible was drilled). This is not a proof that a
bone is modern, for the property of peeling under the shearing action of a rotating
burr is a function of the extent to which the collagen ground-mass of the bone is
preserved ; and under exceptional conditions this has persisted intact even since
Pleistocene times.

Percentages

F N C H2O Ash

Fresh bone . . . .0-03 4-1 14-0 25 53

Piltdown nasal bone . . .0-21 3-9 20-9 27 59

Piltdown " turbinal ". . .0-28 1-7 6-1 25 58

Piltdown I calvaria (maxima) .0-18 1-9 7-5 19 ?6 (min. 62)

Apart from the artificial staining, the Piltdown nasal bones differ from normal
fresh bone only in their fluorine content, which is in excess of that found in recently
buried skeletons except in areas of endemic fluorosis (Bell & Weir, 1949 : 89), or
possibly where the soil has been treated with fluorine-rich phosphate fertilisers.

The great thickness of the Piltdown cranial bones is remarkable (e.g. maximum
thickness of the parietals 12 mm.), but not unique. Cross-sections of the bones show
that the thickness is accounted for by an expansion of the diploe tissues ; the inner
and outer tables are relatively very thin. In all those palaeolithic skulls with very
thick cranial walls that have been examined histologically, the tables are nearly as
thick or even thicker than the diploe (Text-fig. 5). In its structure and thickness
the Piltdown skull can be matched exactly among some recent crania, for example a
skull of an Ona Indian from Tierra del Fuego in the British Museum collection

THE SOLUTION OF THE PILTDOWN PROBLEM

259

(1938.8.10.2). However, such skulls are undoubtedly rare, and to find two in the
same condition at one site would be most unlikely.

The thickening of the diploe in the cranium may be a reflection of a severe chronic
anaemia. The late Professor S. G. Shattock, who examined the Piltdown skull from
the viewpoint of a pathologist reported (1913 : 46) :

TEXT-FIG. 5. Thin sections of parietal and frontal bones of Piltdown and other human
skulls, a, Piltdown I, parietal ; b, Piltdown II, frontal ; c, Ona Indian, parietal ;
d, Swanscombe, parietal ; e, Fontechevade II, parietal ; /, Gibraltar I, parietal ?
Or-e drawn from original specimens ; /based on drawing by Shattock (1913). x 3.
del. D. E. Woodall.

" Certain details of the Piltdown calvaria . . . suggest the possibility of a patho-
logical process having underlain the thickened condition. These are : (i) The
extreme thinness of the tables ; the diploe is closed in on either aspect with the
thinnest compact lamina ; such as can be matched . . . [an ancient Egyptian parietal
is quoted] where the thickening ... is incontestably the residue of a morbid process.
(2) The presence of the elevated patches on the inner surface of certain of the frag-
ments already detailed : in the modern adult skull such fine vascular furrows as
there may have been during growth, on the inner aspect of the tabular bones, have
been smoothly filled in. And (3) to this may be added the synostosis which has here
and there taken place at the sutures although the age of the individual is approxi-
mately only 25 years."

The fragments of the so-called second Piltdown skull have also been artificially
stained, for they contain chromium and show partial alteration to gypsum. They
comprise a small piece of occipital bone and part of a right frontal of unusual thick-
ness. The occipital fragment is not remarkable in thickness or any other morpho-

260 THE SOLUTION OF THE PILTDOWN PROBLEM

logical feature, but its neat rectangular outline suggests that it has been trimmed to
that shape. The frontal fragment is also rectangular as though broken deliberately.
This latter piece could belong to the first cranium, with which it agrees in its excep-
tional thickness and unusual structure, yet in its total composition it appears to
have rather more in common with the occipital fragment grouped with it than with
the occipital or other bones of the first skull.

Percentages

O p. p.m.

N C* CaCO3 P2O6 ! Fe CaSO4 Cr eU3O8

Piltdown I :

Left iron to-parietal (av.). i-i 6-8 3-9 18-7 0-8 6 + 1-5 3

Left temporal . . 0-2 4-8 3-6 23-2 0-8 8 ++ 0-7 i

Right parietal . .1-4 5'3 3*o 19-8 0-8 5 ++ o-o <i

Occipital . . . 0-3 6-8 4-5 20-8 0-7 6 ++ 0-2 2

Piltdown II :

Right frontal . . i-i 4-4 1-5 14-6 0-8 10 ++ <o-i <i

Occipital . . . 0-6 3-9 2-0 13-6 0-2 9 ++ <o-i o

* Carbon in organic fraction.

However, when the analyses are compared, it appears possible that the resem-
blances between the two fragments of the Piltdown II group are not original but
are due to the bones having received the same chemical treatment. The fronto-
parietal bone of the Piltdown I group has been less extensively altered to gypsum
than the others, and it shows the highest radioactivity, the highest chromium
content and the highest carbonate content. The Piltdown II bones, to judge from
their high iron, low carbonate and low phosphate content, received a more intensive
treatment with an acidic iron salt than any of the Piltdown I bones ; and it seems
probable that their lower radioactivity1, lower chromium content and lower carbon
content also reflect differences in treatment.

The feature in the composition of bones which is least likely to be affected by the
iron sulphate and chromate treatment is the fluorine/phosphate ratio, and in this
the frontal of II agrees with the bones of skull I and not with the occipital which
was placed with it.

These more detailed investigations therefore lend further support to the provi-
sional conclusion reached in our 1953 report (p. 145) that the right frontal fragment
originally formed part of the first skull. It is probable that the second skull, from
which the thinner occipital fragment was obtained, was in a sub-fossil condition
similar to that of the first skull.

The human cranial fragments (frontal, parietal and zygomata) reported to have
been found by Dawson in gravel of the Piltdown terrace at Barcombe Mills appear to

1 It has been suggested that the radioactivity of some of the Piltdown bones may be correlated with
traces of K*° due to their having been treated with a solution of potassium dichromate. However,
direct tests failed to demonstrate any exact correlation between the potassium content of the specimens
and either their radioactivity or their chromium content. It is probable that after the bones had been
stained some of the potassium ions would have been removed in solution while the chromium ions became
fixed.

THE SOLUTION OF THE PILTDOWN PROBLEM 261

be pieces of two or possibly three skulls. All these fragments have been artificially
iron-stained, by the sulphate process, but unaided by a chromium compound. In
composition they are broadly comparable with the Piltdown calvaria, but they
differ from all the fragments of these, excepting the occipital of Piltdown II, in
their thickness and structure, which approximate more closely to the normal.

The possibility that occipital II belonged to one of the " Barcombe Mills " skulls
has been considered, but only the parietal agrees in nitrogen content, and this
differs in showing a weak radioactivity which may be original since it cannot be
attributed to K40 in potassium dichromate. In contrast occipital II shows no
radioactivity.

262

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6. CHEMICAL CHANGES IN BONES:
A NOTE ON THE ANALYSES1

By C. F. M. FRYD

Department of the Government Chemist, London

IT is obvious that organic material must undergo changes of a chemical nature during
the process of fossilization, and early in the last century Middleton (1844) pointed
out that some at least of these changes were not complete in periods which were
measurable in geological terms. Carnot (1893) examined the rate of accumulation
of fluorine in bones as a measure of age. Results indicated a general trend, influ-
enced so greatly by local variables as to be of no direct value. However, Oakley
(1948) suggested that the fluorine content of bones from a limited area, as at Piltdown,
or from contiguous sites, could with reservations be used to determine the contem-
poraneity or otherwise of fossils found in physical association ; and since that date
experience has accumulated (Oakley & Montagu 1949, Oakley & Hoskins 1950) on the
extent to which this principle can be used. Further, this principle is of application
whether the change investigated is of an additive nature - such as the incorporation
of dissolved fluorine and possibly uranyl ions in the hydroxyapatite or skeletal bone -
or degradative - such as the loss of fat (Gangl, 1936) or carbon and nitrogen (e.g.
Cook & Heizer, 1952) ; or even sometimes when it is of a partly physical nature
such as the accumulation within the porous structure of the bone of oxides of iron
and other metals, calcium carbonate, or other salts.

In the present investigation we in the Department of the Government Chemist
have endeavoured to assist in solving the Piltdown problem by determining as
accurately as possible on the very small samples available the content of fluorine,
phosphate and iron by adaptation of published methods, and chromium and potas-
sium spectographically. The content of SO4 has also been determined on one sample.
In addition, carbonates have been determined by the use of an apparatus devised
for the purpose (Fryd 1954) ; the carbonate content is of value both as measure of
calcareous infiltration and consequent sealing off of originally porous material, as
an indication of pH in past conditions and as a necessary subtraction from the total
CO2 obtained in the process of determining organic carbon content. Attempts
have been made to determine the fat content of fossil bones by adaptation of pub-
lished methods, but so far no method of value for samples of a very few milligrams in
weight, such as are available from Piltdown material, has emerged. The content of

1 Published by permission of the Government Chemist.

;-3 THE SOLUTION OF THE PILTDOWN PROBLEM 267

organic carbon has been calculated in this laboratory from a combination of our own
figures with those supplied by Drs. Weiler and Strauss of Oxford.

The water of the bone samples was determined by Drs. Weiler and Strauss in
the course of their normal routine of C-H analysis, and not in accordance with the
special procedure adopted by Cook & Heizer (1952) which involves preliminary
drying at 90° and consequently yields lower values.

The nitrogen content of samples was determined in some cases by Mrs. A. Foster
using a modification of the micro-Kjeldahl method devised by her in collaboration
with Dr. J. D. H. Wiseman ; and in other cases by Drs. Weiler and Strauss using
the Dumas method adapted for small samples.

It is hoped to publish full details of the experimental work carried out in the
Government Laboratory elsewhere in the near future (Hoskins & Fryd, 1955).

7. THE X-RAY CRYSTALLOGRAPHY OF THE

PILTDOWN FOSSILS

By G. F. CLARINGBULL AND M. H. HEY

Department of Minerals, British Museum (Natural History)

IN order to evaluate a suggested method of fluorine age determination by measure-
ment of changes in the X-ray diffraction pattern of hydroxy-apatite with variation
of fluorine content (Niggli, Overweel, & van der Vlerk, 1953) a number of bone
samples of varying ages were heated to re-crystallize the apatite. One of these
heated samples gave an unidentifiable diffraction pattern and was thus unsuitable
for the application of the method. This sample, a drilling from the Piltdown cranium,
on examination by X-ray diffraction in the unheated condition, was found to be a
mixture of gypsum and apatite. A subsequent examination by the same method
of many of the Piltdown specimens (see Table VIII) showed that gypsum was widely
distributed in them. Large amounts were found in most specimens except tooth
enamels.

With the object of accounting for this unusual occurrence of gypsum in bone,
not apparently previously recorded, samples of gravel and loam were collected from
the Piltdown site and were examined by chemical and X-ray diffraction methods.
A sample of water from a well £ mile NE. of the site was also examined for sulphate.
The sulphate contents of both water and soil samples, as determined by Dr. Roy C.
Hoather and Dr. C. Bloomfield, are notably low.1 Separation of fine fractions from
the loam and gravel samples failed to disclose the presence of sulphate minerals
and gave only diffraction patterns of clay minerals and quartz. These results tend,
therefore, only to emphasize the anomalous character of the the gypsum in the bone
and to rule out its introduction by natural processes. As there was no obvious
reason for the intentional introduction of gypsum as such into the supposed fossils
it seemed likely that it might have resulted from reaction with a solution used for
colouring more recent bones to simulate iron-stained fossils.

An additional line of investigation supported this theory of chemical treatment.
Part of a skull from Barcombe Mills in the C. Dawson collection at the British
Museum (Nat. Hist.) had some adhering matrix. The skull fragments were found
to have a composition similar to that of Piltdown, namely gypsum and apatite,

1 Water from the well of the " Piltdown Man " Inn, kindly analysed by Dr. Hoather in the Counties
Public Health Laboratory, London, contained 63 parts per million SO8 ; that from Barcombe Mills
47 parts per million. Dr. C. Bloomfield of the Rothampstead Experimental Station found the SO,
in the fine fraction of the Piltdown gravel to be 3-9 mg. per 100 g.

Geol.

Dept.

Reg. No.

X (Film No.

£.590

6346

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6343

£.592

6342

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• 6348

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• 6365

THE SOLUTION OF THE PILTDOWN PROBLEM

TABLE VIII. — X-ray examination of Piltdown specimens

Nature of specimen

Piltdown Skull I, left fronto-parietal

,, ,, I, left temporal

,, ,, I, right parietal

,, ,, I, occipital

,, ,, I, additional fragment of occipital

Piltdown Mandible, bone

,, ,, molar

Mastodon, molar
Elephas cf. planifrons, molar
Hippopotamus, molar

,, premolar

Cervus elaphus, antler

,, ,, metatarsal
Equus, molar
Castor, molar
Piltdown Skull I, nasal

,, ,, I, turbinal

,, Canine

Worked elephant bone
Fragment of bone from base of gravel

269

Estimated
gypsum
content

Castor, incisor

mandible

Elephas cf. planifrons, molar
Mastodon, molar
Rhinoceros, molar
Barcombe Mills Skull, frontal
,, ,, ,, parietal

„ „ zygoma
Piltdown Skull II, frontal
,, ,, II, occipital

„ II, isolated molar
Cervus, tibia

tr.

+

but the matrix was found to contain calcium and ammonium sulphate, although
the fine fractions of freshly coloured gravel from the locality contained no appreciable
sulphate and consisted of clay minerals.

The most obvious method of producing a brown iron stain on bone is by simple
soaking in a solution of a suitable iron salt. Since gypsum is much more soluble in
neutral solutions than is hydroxy-apatite, but is less soluble than the latter in suffi-
ciently acid sulphate solutions, it appeared possible that a fairly acid iron sulphate
solution used for staining might at the same time convert part of the bone to gypsum.
On general chemical grounds, it seemed possible that there might only be a narrow
range of pH within which the solution would be sufficiently acid to convert hydroxy-
apatite to gypsum but not too acid to deposit iron oxide and stain the specimens.
In fact the range of effective pH seems to be quite wide, at least pH 2-5 to pH 6.

270

THE SOLUTION OF THE PILTDOWN PROBLEM

Iron alum (ferric ammonium sulphate) has a distinctly acid reaction and it was
expected that strong solutions might effect some conversion to gypsum. Experiments
showed that even a solution as weak as 2^% was effective in this direction and at'
the same time produced fairly full brown colour. Ferrous sulphate and ferrous
ammonium sulphate are practically neutral in reaction and would not be expected
to produce gypsum so readily, but a few experiments were made with ferrous sul-
phate. These showed that although it had much less reactivity than iron alum
some replacement of apatite by gypsum could be effected. The results of these
experiments are given in Table IX ; Plate 31 shows the X-ray diffraction photo-
graphs.

It seems reasonable to conclude, therefore, that the gypsum in these Piltdown
specimens is the result of their treatment with solutions of an iron sulphate. In
order to produce a similar replacement of the hydroxy-apatite of buried bone it
would appear necessary to postulate a soil composition quite unlike that at Piltdown
or indeed anywhere except perhaps in the close proximity of a sulphide ore-body
undergoing active weathering.

Expt.
Num-
ber

A
G

C

F

M

N
O

TABLE IX. — Artificial iron-staining of bone

Degree of
conversion
of apatite
to gypsum

Solution used Treatment

WHITE BONE, SUB-FOSSIL (NEOLITHIC SKULL, COLDRUM, KENT)

Colour after
treatment

. Saturated iron alum . 36 hr. ca. jo°C.
„ „ .6 hr. ca. jo°C.

. io% iron alum

5%

. Complete . Yellowish-brown
Almost . Yellowish-brown
complete

48 hr. room temp. . Complete . Off white — unchanged
12 hr. room temp. . Partial . Very pale buff, little

change

96 hr. room temp. . Complete . Reddish-brown
then NH3 vapour
for 2 hours
Ditto Partial

2617

. Saturated ferrous . 24 hr. ca. jo°C.

sulphate
. Ditto . 40 hr. room temp.

FRESH BONE
. Saturated iron alum . 9 hr. ca. jo°C.

Light brown
Patchy, reddish-brown
Variable chocolate

brown
Light brown

. Pale yellow — little
change

8. THE BLACK COATING ON THE
PILTDOWN CANINE

By A. E. A. WERNER AND R. J. PLESTERS

Research Laboratory, National Gallery

Optical examination of cross-section

In a cross-section of the black coating adhering to a fragment of the root of the
canine (Text-fig. 6) the following layers can be distinguished :

(a) very thin scattering of a white crystalline material on the surface, thickness
c. 0-015 mm. ; (b) three dark brown structureless layers (the middle one almost
black) probably comprising the cementum layer of the tooth, c. 0*1 mm. ; (c) stained
outer zone of the striated dentine, pale brown semi-transparent, c. 0-2 mm. ; (d) very
thin zone of pale yellowish staining, c 0-015 mm. ; (e) thin zone of orange pigmenta-
tion, which shows under high power a reticulate structure, c. 0-02 mm. ; (/) white,
semi-transparent dentine with characteristic oblique striation.

Chemical examination of cross-section

The zoned appearance of the cross-section suggested that some material from the
black coating had been absorbed preferentially through different depths. The
surface was treated with organic solvent but little or no solvent effect was noted.
It was then treated with dilute hydrochloric acid and a drop of potassium ferro-
cyanide solution. Excess was washed off with distilled water. The top layers
showed a dense mass of Prussian blue, but the pale brown zone below remained
unchanged (except where the precipitate had floated into the cavities). Zone d was
coloured a pale green. Zone e was dark blue and Prussian blue penetrated along the
striations into the unstained dentine (/).

TEXT-FIG. 6. Thin-section of a fragment of the root of the Piltdown canine, showing
the zones of staining, x 30 (approx.).

272 THE SOLUTION OF THE PILTDOWN PROBLEM

Chemical examination of scrapings of the black coating

Under the microscope the scrapings appeared as deep golden brown translucent
lumps ; it was difficult to distinguish between pigment and medium, but a few
reddish brown pigment particles were visible. The usual resin solvents — acetone,
ethyl alcohol, benzene, chloroform — only extracted a small quantity of material
from the sample. The thin cloudy ring of extracted material fluoresced faintly in
ultra-violet light. After evaporation of the solvent, the sample itself was left as a
hard intact mass. (Paint having an oil-resin medium often behaves in this way, a
little of the resin being extracted by organic solvents, yet the paint film itself re-
maining apparently unchanged. Bituminous surface coatings also behave in a
similar manner, a little transparent material being extracted by such solvents).
Aqueous ammonia had a slight solvent effect, and alcoholic ammonia slightly greater.
Morpholine and alcoholic sodium hydroxide softened and gradually disintegrated
the sample. The golden brown transparent material remained dissolved completely
in concentrated hydrochloric acid giving a yellow solution. This solution gave a
copious precipitate of Prussian blue with potassium ferrocyanide, and a strong red
coloration with ammonium thiocyanate. It must therefore contain ferric iron.
From the appearance and solubility of the material this would seem to indicate the
presence of a transparent iron-oxide pigment.

Further scrapings were heated in a small combustion tube. Heavy brownish
fumes with a tarry smell were evolved, and condensed in the form of brown droplets.
This suggested the presence of some bituminous material and would also be con-
sistent with the behaviour of the scrapings to solvents. The incombustible residue
dissolved in concentrated hydrochloric acid, and gave very strong positive reaction
for iron (Fe+++).

Conclusions

The black coating contains a considerable amount of a transparent brown iron-
oxide pigment; it contains some organic matter which seems to be of a bituminous
nature ; and the solubility tests do not exclude the presence of a little oil and /or
resin. It therefore seems that the black coating is a paint consisting of a natural
bituminous pigment, such as Cassel Earth or Cologne Earth (Vandyke brown),
which contains a fairly high proportion of iron oxide, rather than a pure iron oxide
pigment mixed with bitumen. The crackle pattern of the paint and its tough,
non-brittle character (as examined on the tooth itself) are consistent with the above
findings, and would suggest moreover that the surface coating is not of a very great
age.

9. THE GEOLOGY OF THE PILTDOWN
NEIGHBOURHOOD1

By F. H. EDMUNDS

Geological Survey of Great Britain

THE geology of the Piltdown neighbourhood was described by Charles Dawson in
the introduction to his joint paper with Smith Woodward on the Piltdown Skull
(1913 : 117). No geological survey of the superficial deposits of the area had then
been published. In 1925 the writer, in the course of his official work on the Geological
Survey of Great Britain, mapped the superficial deposits of a large area around
Lewes on the scale of 6 in. to the mile. The spread of gravel which according to
Dawson yielded the Piltdown Skull and other fossils was included in this area.

The Piltdown gravel spread rests on the Tunbridge Wells Sand formation. It
measures but 75 yd. x 270 yd. and is one of a number of fragments of a well-defined
but much dissected and denuded river terrace which borders the River Ouse. This
terrace constantly maintains a height of about 50 ft. above the river, and the Pilt-
down spread itself is no exception. The loo-ft. contour of the 6-in. Ordnance
Survey maps, Sussex 40 NW. and NE. (1911 and later editions), actually crosses the
gravel spread, while the 5o-ft. contour almost touches the River Ouse at Gold Bridge,
the nearest point of the river. This is shown on the accompanying sketch-map.

The obvious correlation is with the 5o-ft. terraces of the Thames and other English
rivers ; that is, the gravel is of Upper Pleistocene age.

In 1925 there were some two feet of brickearth lying on an equivalent thickness
of gravel, which rested on Tunbridge Wells Sand. The gravel was seen to be light
brown to orange-red in colour, and to consist largely of sand, with small waterborne
pieces of ferruginous sandstone and ochreous flints. Many of the flints were pitted
with round or pentangular spaces. The brickearth appears to be water deposited,
and to be only slightly later in age than the gravel. It may or may not be contem-
poraneous with extensive brickearth deposits of Upper Pleistocene age in Kent,
some of which have been equated with the loess of Europe. It is of insufficient
importance for separate distinction on the Geological Survey map.

Not unexpectedly the detailed survey of 1925 differs in certain particulars from
Dawson's account of 1913. This is particularly the case in the matter of height of
the gravel spread above the river. Dawson estimated this at about 80 ft. Unfor-
tunately, Sollas (1924 : 183) accepted Dawson's figure without question and equated
the terrace with those at about 100 ft. (or 30 metres) above other rivers of Great
Britain and Europe. Previous attention has been drawn to this false correlation
(Edmunds, 1950).

1 Communicated by permission of the Director, Geological Survey and Museum.

GEOL. II, 6. 22

274

THE SOLUTION OF THE PILTDOWN PROBLEM

The Geological Survey memoir on the Lewes district (White 1926 : 63) repeats
Dawson's estimate in the text. The present writer contributed a text-figure to
this memoir (White 1926, fig. 10) indicating the true elevation of the terrace. The
significance of this diagram, however, does not seem to have been fully appreciated
by Osborne White and the memoir shows a discordance between the text and the
diagram.

Alluvium :

50ft. Grsvel Terrace : '• '•

Tunbridge Wells Sand'

200

Yards v- • '°° — - Contours

TEXT-FIG. 7. Geological sketch map of the Piltdown district.

In his 1913 account, Dawson records that the gravel occurred a few inches beneath
the surface of the soil, and varied in thickness from 3 to 5 ft. In a supplementary
paper (Dawson & Woodward 1914:82) he elaborates his description of the gravel
section, to say that beneath the surface soil there occurs a bed of undisturbed
gravel, from a few inches to 3 ft. in thickness which (he records) overlies a third
bed, which though not always present, is well marked, and consists of pieces of
ironstone and deeply patinated and iron-stained flints.

THE SOLUTION OF THE PILTDOWN PROBLEM 275

The supplementary paper also contains a furthur version of the section in the
form of a drawing annotated as follows :

1. Surface soil, with occasional iron-stained subangular flints . . . Thickness
= i foot.

2. Pale-yellow sandy loam, with small lenticular patches of dark ironstone
gravel and iron-stained subangular flints . . . Thickness = 2 feet 6 inches.

3. Dark-brown ferruginous gravel, with subangular flints and tabular iron-
stone floor covered with depressions. 18 inches.

4. Pale-yellow finely-divided clay and sand, forming a mud reconstructed
from the underlying strata. Certain subangular flints occur, bigger than those
of the overlying beds. Thickness, 8 inches.

5. Undisturbed strata of the Tunbridge Wells Sand (Hastings Beds, Wealden).

(The indicated omissions from the above annotations concern fossils and flint
implements).

This latter description accords reasonably well with the writer's observations
made in 1925, and with a photograph of the section in Dawson's 1913 paper which
shows what is obviously brickearth, about 2 ft. thick, overlying a similar thickness
of gravel.

Traces of the 50 ft. terrace, in the form of remanie flints and sandstone in the surface
soil are visible over much of the countryside bordering the River Ouse. They have
not been noted above the general level of this terrace.

No great significance is to be attached to the mere presence of flints within the
Weald. Dark-stained flints occur at many localities and on present-day views of the
denudation of the Weald their presence is to be expected. Flints occur, however,
in greater numbers in the Piltdown area than have been noted in other districts on
the outcrops of the Weald Clay and earlier formations.

It is reasonable to assume that some older drift deposits may have formerly
existed at higher levels in the Piltdown district. During Pleistocene times, how-
ever, denudation had been so extensive that no field evidence of any earlier drifts
remains. The possible former presence above the 5o-ft. terrace level of a local drift
of any age can only be hypothetical.

io. THE RADIOACTIVITY OF THE
PILTDOWN FOSSILS1

By S. H. U. BOWIE AND C. F. DAVIDSON

Atomic Energy Division, Geological Survey of Great Britain

DURING recent investigations into the geochemical distribution of the radioactive
elements, it was demonstrated by the Atomic Energy Division of the Geological
Survey that fossil bones, teeth and other phosphatic materials tend to accrete
uranium by adsorption from percolating groundwaters (Davidson & Atkin, 1953).
There is some evidence that the radioactivity of a fossil bone is dependent upon
the geological age of the fossil, upon the permeability of the formation in which
it is found, and upon the uranium content of the percolating waters throughout the
ages. In favourable circumstances the determination of radioactivity should there-
fore provide a means of distinguishing older, derived fossils from contemporary
bones when vertebrate remains of more then one age are found in the same geological
environment. No intensive research on this topic has yet been undertaken since
such academic studies are rather far removed from the Atomic Energy Division's
primary function of finding workable uranium deposits.

The build-up of radioactivity in older bones is due to two separate and unrelated
causes — firstly, to the longer time that these fossils have had to adsorb uranium
from circulating waters and, secondly, to the increase in radioactivity as this uranium
approaches secular equilibrium with its daughter elements. The adsorbed uranium
has at first only about one-third of the beta radioactivity of uranium in equilibrium
with its daughter elements, and the content of the latter disintegration products
does not reach a maximum until equilibrium is reached — i.e. until the amount of each
transitory daughter element newly generated in a given time is equal to the amount
lost by decay. If uranium were present in the bone in a known amount when it
was buried, and were not added to or leached away throughout its later history,
the radioactivity would gradually increase for a period of several hundred thousand
years, and the absolute age could be deduced therefrom. Since, however, the
uranium slowly accretes after burial the radioactivity measurements can be no
more than a relative indication of age between different bones found in the same
geological environment.

When The Solution of the Piltdown Problem was published we suggested to the
Keeper of Geology that radioactivity measurements might yield additional informa-
tion on the relative ages of the fossils. He thereupon made a large number of
bones and teeth available for radiometric assays, the results of which are reported
in this note.

1 Communicated by permission of the Director, Geological Survey and Museum.

THE SOLUTION OF THE PILTDOWN PROBLEM 277

The study of phosphatic materials by autoradiographic techniques (Bowie, 1951)
has shown that in radioactive bones, teeth and apatite crystals the radioactivity
may be concentrated towards the surface of the specimen if the latter is relatively
impermeable, but if the material is porous the adsorbed uranium tends to become
evenly distributed throughout. Both for this reason, and because of the high
sensitivity intrinsic to the method, the radiometric analyses of the Piltdown speci-
mens have been conducted by measurement of the beta radiation emitted from the
surface of the bone, this radiation emanating from a layer about 5 mm. in thickness.
The technique employs a sensitive beta counter, with a thin mica end-window placed
at a fixed distance (i cm.) from the fossil under study, the assemblage being contained
in a lead chamber to reduce extraneous radiation to a minimum, and connected to a
scaling unit. One great advantage of the method is that it does not consume any
of the fossil material, which is preserved unchanged. A disadvantage, with the
equipment at present available to us, is that the specimens must be small enough
(less than six inches in length) to permit their introduction into the lead chamber,
and large enough (about a half-inch minimum diameter) to cover the sensitive area
of the end-window counter.

Because of the variation in background count due mainly to fluctuations in cosmic
radiation, the accuracy of the measurements (which is a function of the total
counts recorded) depends upon the length of time allowed for the determinations.
The radioactivity of a relatively uranium-rich bone can be determined with reason-
able precision in a few minutes, but specimens in which the radioactivity is very
feeble demand an investigation lasting one or two days. In the table below a
" standard error " is given, this being a statistical expression of the standard devia-
tion of the background count relative to the count given by the specimen plus
background. It will be observed that in certain instances the standard error
exceeds the count rate, the relevant specimens being devoid of any radioactivity
determinable under these experimental conditions.

From the net count per minute given by each specimen, by extrapolation from
analysed standards, the radioactivity has been expressed as equivalent urania
(£.U3O8). This is a measurement of the true (chemical) uranium content only if
the radioactive elements are in secular equilibrium and if no radioactive elements
other than those of the uranium series are present. In all such phosphatic materials
hitherto analysed chemically, no elements of the thorium series have been found in
significant amounts ; but in the Piltdown specimens which have reputedly been
treated with potassium chromate, part or all of the radioactivity may be due to the
potassium isotope K40 if the potash salts have not been thoroughly washed out from
the bone structure. One per cent. K2O has a beta radioactivity equivalent to about
0-0007% Us^s-

Since we have no personal knowledge of the history or provenance of the Piltdown
and other specimens submitted to us, it is not appropriate that we should attempt
to interpret the results of these tests. There is a strong suggestion that the radio-
activity of the bones varies sympathetically with the fluorine content, particulars
of which have already been published by Oakley ; but since in relatively young
fossil bones the adsorbed uranium cannot have reached secular equilibrium, for such

278 THE SOLUTION OF THE PILTDOWN PROBLEM

materials radioactive measurements may be less satisfactory than fluorine deter-
minations as pointers to age. Possibly, however, radiometric assays may be of
greater value than fluorine determinations for older fossils. It should be noted that
the dentine and cementum of teeth is nearly always more radioactive than the enamel
(PI. 30, fig. 13).

The analyses which we have obtained on a group of late Tertiary and Quaternary
bones, listed below, suggest a rough correlation of radioactivity with age.
Although there are too many variables governing the adsorption of uranium into
such materials for radioactivity measurements in themselves to form a reliable
means of dating the fossils, radiometric determinations seem likely to provide the
palaeontologist with information which, considered in conjunction with other
evidence, may be an important help in discerning the relative age of two or more
groups of vertebrate remains found in the same geological environment.

THE SOLUTION OF THE PILTDOWN PROBLEM

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X

ii. THE FLUORIMETRIC DETERMINATION OF
URANIUM IN THE PILTDOWN FOSSILS1

By A. D. BAYNES-COPE

Department of the Government Chemist, London

IN 1950 Dr. K. P. Oakley asked Dr. C. R. Hoskins of the Department of the Govern-
ment Chemist to investigate the estimation of uranium in fossil bones, for use as a
dating element to supplement the fluorine method.2

Dr. Hoskins' preliminary work showed that chemical methods were not sufficiently
sensitive for the extremely small samples available, and that a sensitive fluorimeter
was necessary. The Chemical Research Laboratory had developed two methods
for the isolation of uranium (see Chemical Methods for the Detection of Uranium,
Stationery Office, London, 1950 ; and U.S. Geol. Surv. Circular No. 199). One,
involving chromatographic separation on a cellulose powder column was found to be
useless for the samples being examined. The other, using solvent extraction into
ethyl acetate, seemed to be adaptable to the estimation of uranium in small samples,
and Mr. R. A. Wells of the Chemical Research Laboratory kindly gave instruction
in the technique used.

The method finally adopted was as follows :

A sample of bone (2 to 10 mg.) was weighed into a Pyrex tube (13 cm. x 075 cm.
I.D., with a B.io stopper) and refluxed with 0-2 ml. of cone, nitric acid until solution
appeared to be complete. After cooling, saturated aluminium nitrate (0-5 ml. of a
solution in i% nitric acid), 0-5 ml. of water and 1-5 ml. of carefully purified ethyl
acetate were added. The mixture was shaken and after settling, i -o ml. of the ethyl
acetate was pipetted on to 0-7 gm. of sodium fluoride. The solvent was evaporated,
the sodium fluoride dried under infra-red lamps, and finally fused. The amount of
uranium was measured in the fluorimeter.

Factors limiting the accuracy of the results were :

(1) Weighing was to the nearest O'i mg. only.

(2) The conversion factor from scale reading to uranium content was em-
pirical.

(3) The lowest readings did not differ enough from the background readings
to admit of satisfactory determination.

(4) Fluctuation in the meter readings allowed only estimation of the second
significant figure.

(5) The carry over of the multiranger was not accurate at the ends of the
scale, and errors of 20% could occur.

1 Published by permission of the Government Chemist.

2 The request was inspired by information received from Dr. C. F. Davidson and Professor Harrison
Brown. K.P.O.

284 THE SOLUTION OF THE PILTDOWN PROBLEM

Taking these factors into consideration only one significant figure can be given
for the uranium percentage.

The uranium adsorbed in bone has been shown by Davidson & Atkin (1953) to
be concentrated in the surface layers and along cracks and crevices, not necessarily
uniformly over any area. A " fair sample " of the bone for analysis must, then, have
a representative proportion of the outer layer to the inner material, if valid
deductions as to the uranium content of the bone as a whole are to be drawn from
the analysis of a single powdered sample.

The results are of value in the following respect : The radioactivity of the Piltdown
fossils could have been due either to uranium or to potassium of isotopic mass 40,
and it can be shown that i% of K2O is equal in ft particle activity to 0-0007% =
7 p.p.m. of uranium. The results obtained are accurate enough to show that the
radiometric assays of Bowie & Davidson give a reliable figure for the uranium content
of the samples examined, though difficulties in sampling, arising from differences
in uranium adsorption with different bone or tooth structures, necessitate very care-
ful interpretation of the results if the correct proportion of uranium in the bone as a
whole is to be estimated.

TABLE XII. — Uranium content of Piltdown and other fossils.

Reg. No. Description % U3O8 U3O8 p.p.m.

£.590 . Piltdown I, left parietal ...... 0-0004 • 4

£.591 . ,, I, left temporal ..... 0-0004 • 4

E.592 . ,, I, right parietal ..... 0-0002 . 2

E.593 . ,, I, occipital . . . . . . 0-0008 . 8

E.594 . ,, mandible ...... <o-oooo2 . <o-2

£.596 . ,, Elephas cf. planifrons molar . . o-i . 1000

E.597 . ,, Elephas cf. planifrons molar, cementum . o-i . 1000

E.62O . ,, Elephas cf. planifrons molar, cementum . o-i . 1000

E.622 . ,, Mastodon molar ..... 0-002 . 20

M.I 5709 . Swanscombe skull, occipital ..... 0-003 • 3°

M. 17034 . Ichkeul "Elephas cf. planifrons" molar, dentine . 0-02 . 200

I acknowledge, with much pleasure, the loan of a fluorimeter by the Atomic
Energy Research Establishment, Harwell, and the help given by Mr. R. A. Wells
of the Chemical Research Laboratory.

12. REFERENCES

ARAMBOURG, C. & ARNOULD, M. 1950. Note sur les fouilles paleontologiques exe'cute'es en

1947-48 et 1949 dans le gisement villafranchien de la Garaet Ichkeul. Bull. Soc. Sci.

nat. Tunis, 2 : 149-157, pis. 17-19.
BELL, G. H. & Weir, J. R. de V. 1949. Physical Properties of Bone in Fluorosis. Industrial

Fluorosis. Med. Res. Council Mem., London, 22 : 85-92.
BOULE, M. 1915. La Pal6ontologie humaine en Angleterre. Anthropologie, Paris, 26 : 1-67,

21 figs.

1923. Les Hommes Fossiles. 2nd ed., xi + 506 pp. Paris. (Esp. pp. 158-176.)

BOWIE, S. H. U. 1951. Autoradiographic techniques in geological research. Bull. Geol.

Surv. G. B., London, 3 : 58-71, pi. 3.
BREUIL, H. 1938. The use of bone implements in the Old Palaeolithic. Antiquity, Gloucester,

12 : 56-67.

• 1949. Silex tailles de Piltdown (Sussex). Bull. Soc. prehist. Fr., Paris, 46 : 344-348.

CARNOT, A. 1893. Recherches sur la composition g6n&rale et la teneur en fluor des os modernes

et des os fossiles de differents ages. Ann. Min. Paris, (9, M6m.) 3 : 155-195.
COOK, S. F. & HEIZER, R. F. 1947. The quantitative investigation of aboriginal sites :

Analyses of human bone. Amer. J. Phys. Anthrop., Washington (n.s.) 5 : 201-220.
• — • — 1952. The fossilization of bone : Organic components and water. Rep. Univ.

Calif. Arch. Surv., Berkeley, 17 : 1-24.
CURWEN, E. C. 1937. The Archaeology of Sussex. 338 pp., 32 pis. London. (Esp. pp.

36-43-)
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EXPLANATION OF PLATES
PLATE 27

Photograph of the medial aspect of (Fig. i) the Piltdown mandible compared with (Fig. 2)
a female orang mandible (unstained) which has been broken in the same manner as the " fossil "
specimen, and in which the first and second molar teeth have been planed down to a correspond-
ing level of dentine exposure. It should be noted that in the Piltdown mandible the angle
has been slightly reduced by abrasion, and the lower margin of the mandibular foramen has
been broken.

[Photographs: Fig. i by C. Horton, Fig. 2 by F. BlackwelL]

Bull. B.M. (N.H.) Geol. 2, 6

PLATE 27

PLATE 28

FIG. 3. (A-C) " Palaeolithic flint implements " recorded from the Piltdown gravel (left to
right £.607, 605, 606) and (D) " Morris's " flint core (£.2690), all showing areas of intense
localised battering. Note that where a chip (arrow) has been removed from E.6o6 the cortex
is pure white below the superficial stain, x 2/3.

FIG. 4. The iron-stained surface of flint £.607 (A above). The speckly character of the
stain is typical of artificial pigmentation of patinated flint. x 2.

FIGS. 5-6. Flaked surface of flint £.607 (Fig. 5) compared with the naturally iron-stained
surface of a flint (£.965) from the Piltdown gravel (Fig. 6). The ink circles indicate areas
which were wiped immediately after dilute hydrochloric acid had been applied to them. The
stain on £.607 was removed, the natural stain on £.965 was unaffected. Natural size.

[Photographs : C. Horton.]

Bull. B.M. (N.H.) Genl. 2, 6

PLATE 28

PLATE 29

FIGS. 7, 8. The pointed end oi the Piltdown bone " implement ", showing details of the cuts
on the lateral faces. £.615. Natural size.

FIG. 9. The cut facets at the rounded butt end of the " implement ". Natural size.

FIG. 10. A piece of fossil bone from the Swanscombe gravels which has been shaped with a
steel razor and artificially iron-stained ; it reproduces all the essential features of the Piltdown
bone " implement ". £.2707. x 2/3.

[Photographs : C. Norton],

Hull. li.M. (NJL) Gcol. 2, 6

PL ATM 29

IO

PLATE 30

FIG. ii. Electron-micrograph of decalcified residue of a sample of the Piltdown mandible,
showing banded collagen fibres, x 30,000 (approx.)

Fio. 12. Section of fragment of molar tooth of Elephas cf. planifrons in the Piltdown collec-
tion (£.620). x 2.

FIG. 13. Autoradiograph (intensified) of the same section, produced by six weeks contact
with very sensitive film (Ilford nuclear research pi., emulsion type B.a) ; demonstrating the
high radioactivity of the dentine and cementum layers. x 2.

[Photographs: Fig. n by A. V. W. Martin, Fig. 12 by C. Horton, Fig. 13 by S. H. V. Bowie.]

Bull. B.M. (X.H.) Geul. 2, 6

PLATE 30

13

PLATE 31

X-ray diffraction photographs on 6 cm. diameter camera ; Co-Koc radiation. A. Gypsum
(Mosul marble). B. Apatite (Jumilla, Spain), c. Apatite (Fragment of bone). D. Gypsum
-{-apatite (Piltdown skull, frontal). K. Gypsum (Neolithic skull bone from Coldrum, Kent,
soaked lor 96 hours in 10% solution of iron alum).

[Photographs taken in the Department of Minerals.]

Hull. li.M. (\.H.) Gcol. 2, 0

PLATE 31

PRESENTED

26 JAN 1355

PRINTED IN GREAT BRITAIN BY
ADLARD AND SON, LIMITED
BARTHOLOMEW PRESS, DORKING

1 OCT 1955

THE SCHLZAEACEAE OF

SOUr OF ENGLAND N

EARLY ERTtARY MES

M. E. J. CHANDLER

BULLETIN OF

THE BRITISH MUSEUM (NATURAL HISTORY)
GEOLOGY Vol. 2 No. 7

LONDON : 1955

BULLETIN OF
THE BRITISH MUSEUM (NATURAL HISTORY)

GEOLOGY

The following papers appeared in Volume i (1949-52) :

Price

No. i (1949). The Pterobranch Rhabdopleura in the English Eocene.

H. D. Thomas & A. G. Davis • 1s- 6d-

No. 2 (1949). A Reconsideration of the Galley Hill Skeleton. K. P.

Oakley & M. F. Ashley Montagu . 5*-

No. 3 (1950). The Vertebrate Faunas of the Lower Old Red Sandstone
of the Welsh Borders. E. I. White.

Pteraspis leathensis White a Dittonian Zone-Fossil. E. I.
White
No. 4 (1950). A New Tithonian Ammonoid Fauna from Kurdistan,

Northern Iraq. L. F. Spath .
No. 5 (1951). Cretaceous and Eocene Peduncles of the Cirripede Euscal-

pellum. T. H. Withers . • 5«-

No. 6 (1951). Some Jurassic and Cretaceous Crabs (Prosoponidae) .

T. H. Withers . 5$-

No. 7 (1952). A New Trochiliscus (Charophyta) from the Downtonian

of Podolia. W. N. Croft
No. 8 (1952). Cretaceous and Tertiary Foraminifera from the Middle

East. T. F. Grimsdale.

No. 9 (1952). Australian Arthrodires. E. I. White . ^5S>

No. 10 (1952). Cyclopygid Trilobites from Girvan. W. F. Whittard . 6s.

THE SCHIZAEACEAE OF THE SOUTH OF
ENGLAND IN EARLY TERTIARY TIMES

BY

MARJORIE E. J. CHANDLER

Pp. 291-314 ; Pis. 32-38 ; 2 Text-figures

BULLETIN OF

THE BRITISH MUSEUM (NATURAL HISTORY)
GEOLOGY Vol. 2 No. 7

LONDON: 1955

THE BULLETIN OF THE BRITISH MUSEUM
(NATURAL HISTORY), instituted in 1949, is
issued in five series, corresponding to the Departments
of the Museum, and an Historical series.

Parts will appear at irregular intervals as they become
ready. Volumes will contain about three or four
hundred pages, and will not necessarily be completed
within one calendar year.

This paper is Vol. 2, No. 7 of the Geological series.

PRINTED BY ORDER OF THE TRUSTEES OF
THE BRITISH MUSEUM

Issued September, 1955 Price Fifteen Shillings

THE SCHIZAEACEAE OF THE SOUTH OF
ENGLAND IN EARLY TERTIARY TIMES

By M. E. J. CHANDLER

SYNOPSIS

The paper records the discovery of two new species of Anemia, A . poolensis and A . colwellensis,
and one of Lygodium, L. poolensis, in the early Tertiary strata of Britain. It also gives new
information about Lygodium kaulfussi Heer from fertile material. Fertile pinnules of all these
species are so beautifully preserved that more is now known in detail of some of these fossils
than of many living species. Previous knowledge of British Tertiary Schizaeaceae is
summarized. Fuller and more satisfactory evidence that the two genera once grew in southern
England supports other evidence from fruits and seeds (awaiting publication) of a very warm
and humid climate in the south of England throughout the early Tertiary period.

INTRODUCTION

PREVIOUS knowledge of the Schizaeaceae in the English Tertiary was based on sterile
fronds of Anemia subcretacea (Sap.), Lygodium prestwichii Gard. & Ett. and Lygodium
kaulfussi Hr. together with some imperfectly studied fertile fragments of the last
species. Like the contemporary angiosperms these ferns indicate a very warm and
humid climate. They are perhaps survivors of an ancient widespread more or less
uniform broad Mesozoic tropical belt of vegetation. For Anemia appears to be a
relict genus in the old world, where it is now represented by few species although the
genus was formerly widespread. But it flourishes and is represented by numerous
species in the tropics and subtropics of the New World, albeit with a restricted
northern range. Lygodium still holds its own in both hemispheres with a shrunken
latitudinal area of distribution which can be attributed to the universal post-Eocene
cooling of climate and consequent contraction of the tropical and subtropical belt
of vegetation.

This paper is a bye-product of research upon the fruits and seeds of Tertiary Beds
in Dorset, Hampshire, and the Isle of Wight. The material first to be described
was found in abundance while sifting carbonaceous sands in the Lower Bagshot
Beds (Cuisian ?) on the north shores of Poole Harbour at Lake, near Hamworthy,
Dorset. The matrix was coarse and rich in seams of wood and fruits among which
were small, often crushed and contorted, ovoid or subglobular, segmented bodies.
The majority had tightly enrolled segments of distinctive appearance. When mois-
tened with alcohol or nitric acid on a glass slide and examined under the microscope,
quantities of large globular-tetrahedral spores were released among which were
many larger turgid bolster-shaped or fusiform objects of spore-like appearance.

GBOL. n, 7. 23

292 THE SCHIZAEACEAE OF THE SOUTH OF ENGLAND

Further investigation showed that on the inner surface of the enrolled segments
sporangia were seated apparently in two rows. They had the typical apical annulus
of Schizaeaceae, but the occurrence of what appeared to be two types of spore
presented a problem for which botanists who were consulted could produce no ready
solution.

In due course the specimens were shown to the late W. N. Croft. With charac-
teristic generosity and interest he gave up many hours to the investigation of the
matter, studying with painstaking thoroughness not only the evidence yielded by
the fossils, but all that could be discovered from living material of Anemia. In
addition he searched the literature of Recent ferns, and such published work as is
available on fossil Schizaeaceae in the hope of obtaining a clue to the mystery of the
two types of " spore". ' As the result of a very large amount of work Mr. Croft
discovered that the turgid elongate "spores" were really paraphyses such as are
found growing among the sporangia on the surface of Recent fertile Anemia pinnules.
The relationship of the fossils to Anemia was then demonstrated by cell-structure,
sporangia and spore-characters and a spore-count. Mr. Croft was also responsible
for most of the fine series of highly magnified photographs of sporangia, spores and
paraphyses shown in the accompanying plates.

It was our original intention to produce a joint paper on this species, here named
A. poolensis. But unfortunately Mr. Croft died before the work was finished. It
therefore remains for me to record the results of our efforts and to pay tribute to the
work of a most generous and able colleague. It can be claimed that more is known
of the fruiting organs of the fossil species A. poolensis than of many of the living
species of this large tropical and subtropical genus. The same species was found
represented by one pinnule in somewhat younger beds at Sandbanks and at Branksome
Dene in the Bournemouth Freshwater Beds (Lutetian ?) where several macerated
pinnules occurred. It was represented at Cliff End near Mudeford (Auversian ?)
by a minute fragment which yielded spores, and by a solitary spore at Alum Bay,
Isle of Wight (Lower Bagshot, just above the pipe clay) where, according to Gardner
& Ettingshausen (1880 : 47), Heer was confident he had seen a barren pinnule of
A. subcretacea.

While working on A. poolensis, I had the good fortune to discover a tiny fertile
pinnule of a second species of Anemia in the Upper Headon Beds of Col well Bay,
Isle of Wight. It is here described as A. colwellensis. In spite of prolonged search
and sifting no second specimen has yet been found. It occurred in a small pocket
of broken and battered fragments which included scraps of Gleichenia.

A second genus of Schizaeaceae, Lygodium, represented by fruiting pinnules of
two species also turned up at about the same time. One was closely associated with
barren pinnules of L. kaulfussi at Studland (Cuisian ?) and may be presumed to belong
to that species. It was also found in the Bournemouth Freshwater Beds at Brank-
some Dene, Dorset. The species formerly occurred in considerable abundance at
Bournemouth, chiefly as barren pinnules but rarely as fertile ones. The other was
represented by a naturally macerated fertile pinnule in fine sediment at the base of
the section at Lake. Its spores demonstrated beyond question that it is
specifically distinct from L, kaidfussi. It has been named Lygodium poolensis, A

THE SCHIZAEACEAE OF THE SOUTH OF ENGLAND 293

solitary spore was subsequently recognized entrapped among the hairs of Anemia
on slide ¥.31511.

The importance of these discoveries lies in the fact that hitherto the occurrence
of Schizaeaceae in the south of England has been based on barren pinnules only, if
we except the imperfectly investigated fertile tufts of Lygodium figured by Gardner
& Ettingshausen from Bournemouth (see below). The fertile fragments hereafter
described confirm the earlier work by providing fuller and more satisfactory evidence
that the genera Anemia and Lygodium once grew in southern England. The presence
of this essentially tropical family supports evidence from fruits and seeds (awaiting
publication) of a very warm and humid climate in the south of England throughout
the earlier Tertiary period.

Previous work takes account of three species only : Anemia subcretacea (Saporta)
from the Bournemouth Freshwater Beds and Reading Beds, Lygodium prestwichii
Card. & Ett. from the Woolwich and Reading Beds, and Lygodium kaulfussi Heer
from the Bournemouth and Studland Beds. A brief account of these three species
as recorded in literature is given here, while L. prestwichii is also described from a
new site at Newington, Kent and L. kaulfussi from fresh barren material from Stud-
land as well as from the newly discovered fertile material. A few barren pinnules
from the Bembridge Beds originally described as Filix incertae sedis, sp. 5, can now
be referred with reasonable certainty to a species of Anemia although few of its
characters are known. Its nervation and pinnule margins are strongly reminiscent
of the fertile pinnule segments of A. colwellensis. As there is no definite connecting
link between the two, however, the Bembridge pinnules must be regarded, for the
time being at least, as representing a third species.

Genus ANEMIA Swartz

NOTE : The spelling of the generic name is that given in the code of Botanical
Nomenclature 1952, p. 83.

Anemia subcretacea (Saporta)

1868. Asplenium subcretaceum Saporta, p. 315, pi. 23, fig. 4.

1880. Anemia subcretacea (Saporta) Gardner & Ettingshausen, p. 45, pis. 8, 9.

1882. Anemia subcretacea (Saporta) : Gardner & Ettinghausen, p. 67.

1886. Anemia subcretacea (Saporta) : Gardner, pp. 400, 402, pi. i.

The fern is illustrated and fully described by Gardner & Ettingshausen (1880) from
the Bournemouth Freshwater Beds, and is illustrated by Gardner (1886), but with-
out full description, from the Reading Beds of Reading.

The specimens are preserved in the British Museum (Natural History) Geology
Dept., Nos. ¥.14970-73 and ¥.15033-42 (Bournemouth) and ¥.15272 (Reading).

According to Gardner the barren pinnules were very abundant at Bournemouth
especially to the east of Bournemouth pier, decreasing in size and abundance west-
wards towards Poole Harbour, while at Branksea only small pinnules occurred. He

294 THE SCHIZAEACEAE OF THE SOUTH OF ENGLAND

comments that the fertile pinnules, which he had never seen, were almost certainly
separate and therefore rarely if ever preserved. He added that although the fern
" did not agree very closely with any existing species of Anemia, the general form
and dichotomy of the frond, the venation, the length and strength of the stipes, the
separation of the fertile and barren pinnae, have, after much consideration, induced
us to place it in the genus Anemia". He compared it with A. adiantifolia Sw. from
Cuba, the Bahamas, Florida and Mexico to Bahia.

Gardner & Ettingshausen (1880 : 47) report that Saporta, who had evidently
compared Bournemouth specimens with similar ferns from Sezanne, had no hesita-
tion in stating that the two were identical, while Lesquereux compared them with
some from the Eocene of North America and could see no difference whatsoever.
Gardner & Ettingshausen (1882 : 67) add that the species is characteristic of the
" older Eocene, and even pre-Eocene Rocks". They state that it first appeared in
the Cretaceous and its upper limit in Europe is the Bournemouth horizon and that
it attained its greatest luxuriance in the Eocene Beds of what are now temperate
latitudes although it ranged from the south of France to the Arctic regions.

This barren foliage closely resembles specimens of Anemia fremonti Knowlton and
Anemia sp. figured by Andrews & Pearsall (1941, pi. 5, figs. 30-32, 34, 35) from the
Upper Cretaceous Frontier Formation of south-western Wyoming, U.S.A.

The discovery of abundant fertile pinnules of Anemia, described below as A.
poolensis, in the Bournemouth Beds (Cuisian or Lutetian ?) of the western end of the
Bournemouth section raises the question whether these detached fragments are the
fertile form of A. subcretacea. The relationship cannot be proved in the absence of
any organic connection and in the absence of the two forms in close association in
the same layers. They are separated therefore for the time being although it is
quite possible that future evidence may make it necessary to unite them under one
specific name. Should this eventually happen the character of the spores (p. 301,
PI. 33, figs. 24-31 ; PI. 36, figs. 54-58) would clearly separate A. subcretacea from
A . fremonti forma fertilis Andrews and therefore presumably from the barren A .
fremonti Knowlton. Nevertheless it is interesting that species of Anemia with the
same type of foliage existed in Cretaceous and Eocene times in the two hemispheres,
and that in both the genus extended far to the north of its main present day range in
tropical and subtropical latitudes. In America Anemia is found only as far north as
the Sierra Nevada and Florida but is richly represented by many species in Central
and South America and the West Indies, especially in mountainous regions. In the
Old World today it survives only as a relict genus with a very few species, one A.
lanipes Christensen in Madagascar, one A. schimperiana Presl. in tropical Africa and
South India, and one A . dregeana Kze in Natal. The genus may perhaps be regarded
as a representative of that ancient more or less uniform tropical plant belt of low
latitudes in Cretaceous times. For unknown reasons it persists as a vigorous com-
ponent of the tropical and subtropical flora of the New World up to the present,
but in spite of its former wide range in the old world it remains there now only in an
attenuated form although in the equatorial belt the climate was probably always
favourable to it. But outside the tropics where colder conditions prevailed after the
Eocene it was exterminated altogether.

THE SCHIZAEACEAE OF THE SOUTH OF ENGLAND 295

Anemia poolensis n. sp.

(PI. 32, figs, i-io ; PI. 33, figs. 14-22, 24-31 ; PL 34 ; PI. 35, fig. 41 ; P.I 36, figs.

54-58 ; Text-figs, i, 2)

DIAGNOSIS. Fertile pinnules deeply divided into a varying number of segments,
4 to ? 7, some of the segments being further dissected and often somewhat twisted.
Segments strongly recurved so as to form a compact body about 2 or 3 mm. across
(often open below), foliose, irregularly and deeply toothed or jagged especially
distally, often sharply pointed, sometimes boat-shaped, occasionally clothed with a
felt of pointed usually non-septate hairs which may occur on both surfaces. Spor-
angia numerous, sub-ovoid to sub-globular, about 0-25 to 0-44 mm. long and 0-2
to 0-3 mm. broad. Annulus symmetrical, apical, occupying about two-fifths of the
length of the sporangium, formed of a single row of about fifteen straight thick-
walled cells. Cells of sporangium wall almost straight superficially but when exam-
ined with J in. objective having a finely sinuous appearance. Apical plate delicate
of about 12-14 cells. Trilete tetrahedral-globose spores entirely smooth, about 50 to
6o/* (rarely larger or smaller) in diameter, wall 2 to 2 -5^ thick as seen in optical section.
Paraphyses many among the sporangia, sessile or shortly stalked, about 72 to 156/4
long, 48 to ii4/* broad.

HOLOTYPE. ¥.31484.

DESCRIPTION. Rhizome and frond including fertile stipe unknown. Fertile spike
probably somewhat flattened in one plane, by inference (see below).

Fertile tertiary pinnae having several segmented pinnules of which one is terminal
while the others may spring from one stipe and form a pair of sub-globular bodies
(Text-figs, i c, G, H) or they may arise singly (Text-figs. IE, F ; 2B).

Fertile pinnules divided sympodially into from 4 to at least 7 segments, some of
which may be further dissected and much twisted. Segments strongly recurved
so as to form a compact sub-ovoid, sub-globular, or ellipsoid " body " about 2 to 3
mm. across but varying in size with the degree of folding and recurving, largest in
the least recurved specimens. Terminal pinnules more simply reflexed than the
others in the few such specimens seen, and not enrolled at the distal ends of the seg-
ments so that the sporangia on their lower surfaces are partially exposed (Text-figs.
IA, B, D ; 2G, H), other pinnules are usually open below to a lesser extent, for not all
segments are enrolled distally, so that the " bodies " may gape downwards (Text-figs.
IH ; 2A, E, F). Segments rather rigid, foliose, of variable width without indusium-
like margins, irregularly and sometimes deeply toothed with sharply pointed narrow
teeth and free tips. Hence their distal ends have a jagged appearance (Text-figs.
IA, B, G, H ; 20). These tertiary pinnules (" bodies ") are often somewhat flattened.
This appears to have been an original character as indicated by the form of the seg-
ments, those which form their lateral margins being frequently folded longitudinally
and more or less inverted boat-shaped so that they embrace the adjacent segments
(Text-figs. 1 1, j) . Those over the middle part of the bodies may be broad or unfolded,
they may even be slightly concave outwards with outward curving edges. The boat-
shaped lateral segments are unequal sided as a rule, one side of the " boat " being

296

THE SCHIZAEACEAE OF THE SOUTH OF ENGLAND

TEXT-FIGURE 1

THE SCHIZAEACEAE OF THE SOUTH OF ENGLAND 297

TEXT-FIG, i. Anemia poolensis n. sp.

(A) Side view of a terminal pinnule from a fertile tertiary pinna. Stipe to right. Five
recurved toothed segments (a, b, c, d, /,) are shown with a few sporangia exposed in the gaps
between them. Direction of cell-structure indicated on the upper surface of the segment.
Length, 3-1 mm. x 20 approx. Holotype ¥.31484.

(B) Under surface of the same. More of the recurved segments are seen a, b, c, d, f, as in A,
e, g, additional segments not visible in A. Note the sporangia. Paraphyses were attached
at p. x 20 approx.

(c) A pair of tightly reflexed downwardly directed pinnules (i) and (2) showing the broken
end of the stipe (st) from which they spring. Length, 2-4 mm.; breadth across the pair,
3-25 mm. Surface much sand-pitted in places, x 17 approx. V. 31485.

(D) A terminal loosely recurved pinnule. It was originally united on the left to a stipe which
bore two other fertile pinnules (now detached). Length about 2-5 mm. ; height, r6 mm.
(n) is a nervule flanked by fragments of reduced fronds, (c) Indicates shining cells with
digitate walls. Under the microscope the walls appear raised and rounded, the centre of
the cells depressed. There are sporangia on the lower surface, x 17 approx. ¥.31486.

(E) A tightly recurved and enrolled single pinnule still attached to a large fragment (1-4 mm.
long) of stipe (st) of the tertiary pinna. The midrib of the stipe is channelled on the upper
surface at (v), the channel marking a vein flanked by reduced lamina, (b) and (c) are two
segments. Maximum diameter at right angles to stipe about 2^75 mm. x 15 approx.
V. 31487-

(F) Reverse side of E. (st) stipe, (b, c) correspond to segments so labelled in E. A mass of
spores liberated from a disrupted sporangium is indicated on the right at (sp).

(G) A pair of tightly reflexed downwardly directed pinnules still attached to the remains of a
stipe (st). Length about 2-75 mm.; across the pair, 3 mm. x 17 approx. ¥.31488.

(H) Reverse side of the same specimen showing the reflexed segments of the two pinnules, with

a few sporangia (sp) exposed,
(i) A detached typical unequal-sided boat-shaped segment showing the upper surface. A

sporangia projects from below at (sp) with open stomium. Oblique striae diverge from

the nervule over the surface on this side of the segment. Proximal end to right. Length

preserved 1-8 mm.; depth, 0-75 mm. x 20 approx. ¥.31489.
(j) Opposite side of the same segment which displays more of the under surface owing to the

inequality of the sides of the segment. Proximal end to left. Six sporangia are shown,

the two at the distal end with gaping stomium (sm) directed to the margins of the segment.

Orientation of cells indicated on the upper surface, x 20 approx.

All the above are from Lake, nr. Hamworthy, Dorset.

GEOL. II, 7.

2Q8

THE SCHIZAEACEAE OF THE SOUTH OF ENGLAND

TEXT-FIGURE 2

THE SCHIZAEACEAE OF THE SOUTH OF ENGLAND 299

TEXT-FIG. 2. A-H. Anemia poolensis n. sp. i. Anemia colwellensis n. sp.

(A) Sharply reflexed pinnule with downwardly directed segments (a-/). A few sporangia are
exposed where they gape. The direction of the cells, transverse over the middle of the
segments, diverging near the edges, is indicated. Length, 2-25 mm.; breadth, 2-5 mm.
(g) is the lower surface of the stipe, x 20 approx. ¥.31490.

(B) The reverse of A. Segments and stipe lettered to correspond with A.

(c) Part of an unrolled pinnule, lower surface. Tips of segments at the base of the figure
appear to be broken.. The segments above retain their sporangia (sp). The segment
below shows hollows (h) from which sporangia have fallen. The nervules of the segments
are indicated where they could be traced. Maximum diameter of the specimen, 2-5 mm.
Maximum length preserved measured along the nervule, about 1-5 mm. x 27 approx.
¥.31491.

(D) A pair of pinnules, one large (i) the other small (2) borne on the stipe (st). Segments much
folded together. Surface covered with a felt of hairs, indicated only, in the drawing.
Length, 3 mm.; breadth, 2-75 mm. This specimen yielded a smooth spore identical in
all its characters with those of the smooth hairless specimens (see ¥.31506). x 17 approx.
¥.31492.

(E) A pinnule originally more or less complete but now with the tips of the segments broken.
Segment lettered (d) may be the stipe. Direction of cells indicated on the surface. Spor-
angia and released spore masses are seen in the opening below. Length, 1-5 mm.; breadth,
1-75 mm. x 24 approx. ¥.31493.

(F) Opposite side of the above showing segments (a) and (e) much sand pitted, (d) again
may be the stipe.

(G) Upper surface probably of a terminal pinnule. Sporangia are seen in profile in the segment
at the top of the figure. The stomium opens outwards. Surface much sand pitted.
Maximum length from tip of free segment to base of stipe (?), 2-5 mm.; breadth, 1-75 mm.
X 24 approx. ¥. 31494.

Opposite side of specimen showing the lower surface of the pinnule with a few folded and
twisted segments. Sporangia are seen, but over most of this surface they have been
disrupted so that the surface is concealed by a mass of released spores (s) . x 24 approx.
Lower surface of a fertile pinnule (the only specimen) which yielded all sporangia and spores
figured (based on a drawing made before the specimen collapsed). It shows alternating,
smooth-edged, rounded segments on opposite sides of the nerve or midrib, their recurved
margins (r, r) which produce the smooth outlines form a fringe-like indusium covering
part of the lobes on this surface. The sinuous midrib, and the pocket-like depressions
(p, p) from which sporangia have fallen can be seen. The arrangement of sporangia (sp,
sp) in the lobes on each side of the midrib is indicated, x 20 approx. Holotype. ¥.31522.
Colwell Bay, Isle of Wight.

Figs. A-D, G, H are from Lake, Dorset. Figs. E, F from Sandbanks, Dorset.

300 THE SCHIZAEACEAE OF THE SOUTH OF ENGLAND

considerably broader than the other (Text-figs, i I, j). The flattening of the tertiary
pinnules suggests that the fertile spike had a tendency to be flattened in one plane in
life, but the character seems not to have been constant.

Both surfaces are sometimes clothed with a close felt of red or black pointed usually
single-celled hairs (PI. 32, fig. 6 ; Text-fig. 2D) pitted or corroded in fossilization. In
the majority of specimens seen, these have disappeared, if they were ever present,
and a highly characteristic glistening outer (upper) surface is exposed. This is
formed of transversely elongate cells over the median longitudinal part of each
segment, the cells being also aligned in longitudinal rows. As seen by reflected light
they are rather coarse with sinuous outlines having sunk centres and raised margins
so that they produce a conspicuous and highly characteristic pattern. In section
(PI. 34, figs. 32, 34, 37) more than one layer of these cells is seen and they appear
thin-walled. Towards the edges of the segments the coarse cells give place to finer
narrow elongate ones which form oblique striae diverging from the median band of
shining cells. Inner or lower surface of the pinnule with similar narrow cells which
diverge obliquely from the median line or venule of each segment towards its margins.
Venules with ? scalariform thickening (PI. 34, fig. 36). Granules like chloroplasts
are common in many of the cells and also amongst macerated spores. They may
occur singly or in groups (PL 32, fig. 10 ; PI. 34, figs. 37, 38 ; PI. 35, fig. 41) and appear
sometimes to be aggregated into small irregularly shaped plates in which the boun-
daries between the grains are more or less obscure. The grains are usually rounded
or oval but sometimes subangular. Their average size is about 2-5 to 3'5/*. Similar
granules occur in living Anemia, e.g. in A. phyllitides (L.) where they are greenish
and about 3 -5/4 in diameter but they may be as large as 6fi. In form and occurrence
they are like those of the fossil.

Paraphyses (inflated short sterile filaments) (PI. 33, figs. 18-22 ; PL 34, fig. 37)
are abundant all over the lower surface of the pinnules growing amongst the sporangia.
They are sub-ovoid to ellipsoid, usually straight, rarely slightly curved, sometimes
with a fairly prominent point at the distal end, and a very short stalk at the
proximal end. Sometimes they are asymmetrically attached. A few may be sessile.
When freshly released from the pinnule segments and scarcely macerated they are
shining and turgid, and dark brown in colour. On maceration they quickly collapse,
the brown layer within cracks and tends to disintegrate revealing that the paraphysis
wall is extremely thin in optical section. The size varies from about 72 by 48/4 to
156 by 78/4 or they may be as broad and short as 114 by 114/1.

The Sporangia (PL 32, figs. 4, 5, 7-10 ; PL 33, figs. 14-17 ; PL 34, figs. 33-35, 37 ;
PL 35, fig. 41) are shortly stalked (? sometimes sessile) borne in two rows which are
closely spaced, alternating and forwardly directed. They arise from the venule or
from points close beside it (PL 32, fig. 4). In the inverted boat-shaped lateral
pinnule segments they project conspicuously above the narrow side of the segment
(Text-fig, ij). They are sub-ovoid, rather broad at the base, attached asymmetric-
ally (PL 32, figs. 7, 8). The annulus is apical, symmetrical, well-developed and con-
spicuous, consisting of a single row of about 15 straight-sided cells (excluding the
stomium) the internal and radial walls being strongly thickened. Length of annulus
about two-fifths of the length of the sporangium (PL 32, figs. 4, 5, 7-10 : PL 33,

THE SCHIZAEACEAE OF THE SOUTH OF ENGLAND 301

figs. 14-17). The stomium consists of two to three rows of cells, dehiscence is
longitudinal extending the whole length of the sporangium, normally occurring on
the face towards the outer side of the segments (PL 32, fig. 5 ; PL 33, figs. 14, 15 ;
Text-fig, ij). Apical plate small and circular, very delicate, its distal face with
about 12-14 cells, those nearest the centre of the plate equiaxial, five or six-sided
with straight sides, those near the edge of the plate broader in a tangential than in a
radial direction (PL 33, figs. 15, 17). The cells of the sporangial wall appear straight
superficially with a i-in. or |-in. lens, sometimes they are rounded, at other times
pointed at their extremities, elongate parallel to the major axis of the sporangium.
When seen from the interior or examined with a J-in. lens they show a bead-like
thickening (PL 32, fig. 8) which produces the effect of fine sinuosities on the external
surface. Spore output probably 128 (115 actually counted). Length of sporangium
about 0-25 to 0-44 mm., breadth about 0-2 to 0-3 mm.

Spores (PL 33, figs. 24-31 ; PL 36, figs. 54-58) trilete, tetrahedral-globose,
non-striate, smooth, glistening, normally about 50 to 6o/* in diameter but occas-
ionally as small as 367* and as large as 70-4/4 ; about 2 to 2-5 fi thick, splitting occa-
sionally along the triradiate mark, the distal ends of which sometimes show a fine
forking in unsplit specimens (PL 33, fig. 27 ; PL 36, fig. 54).

REMARKS. These curiously rolled fertile pinnules were chiefly found in the coarser
carbonaceous seams at Lake. Numerous specimens have so far been collected
and many more would reward persistent collecting and sifting. More than sixty
are available for study by reflected light and will be catalogued in a forthcoming
British Museum monograph. Many others have been macerated to varying degrees
and mounted on slides for examination by transmitted light.

The arrangement of the segments was at first very difficult to discover, but once
seen it could be detected almost invariably especially after microscopic examination.
In order the better to show structures which are even more obscure in the photo-
graphs than in the specimens themselves, sketches have been made (Text-figs, i
and 2) which are essential to the understanding of these confusing specimens.

The technique adopted in the study of the material was as follows : they were at
first examined as a whole by reflected light, a method which best showed the form
and alignment of the surface cells, and the arrangement of sporangia and paraphyses
on the segments. Many specimens or segments of pinnules were then macerated
in nitric acid and mounted, some in xylol, others in Euparal or Canada balsam.
They were then examined by transmitted light, and in the case of the denser speci-
mens by strong reflected light using a white background. In this way some of the
most delicate structures were seen and photographed. The apical plates of the
sporangia were most difficult to display as their delicate cells were peculiarly liable
to collapse and tear on maceration. They were most easily seen when mounted in
glycerine ; when transferred to alcohol they were frequently disrupted.

Spores and paraphyses were readily released from the sporangia or pinnule
segments on moistening with alcohol or nitric acid. The spores were tough and
resistant but the paraphyses were so thin-walled that they frequently collapsed as
described on p. 300.

A series of microtome sections were prepared under the direction of W. N. Croft.

302 THE SCHIZAEACEAE OF THE SOUTH OF ENGLAND

His notes show that they were cut by Mr. S. Prudhoe from a specimen which was
thereby destroyed ; it is now represented by slides ¥.31506-07, ¥.31509-10. The
" body " was impregnated with paraffin wax after treatment with hydrofluoric acid
by the method described by Lang (1929 : 667-8). A first series of microtome
sections cut at io/t (slide ¥.31507) produced rather incoherent sections and the speci-
men was re-embedded. A second series was cut at 6/t (slide ¥.31510). Although
less incoherent it was re-embedded. A third series was cut at io/* (slide ¥.31509) . It
held together fairly well but the centre came away as loose powder, due, it was thought
to the toughness of some of the tissues. The remaining piece was softened by treat-
ment with tribasic sodium. The fourth series (slide ¥.31506) was cut from the
softened tissue but it appeared to be little better. It was spoilt by the cover-slip
sliding over the sections during cleaning. Some of the sections are illustrated in
PI. 34, figs. 32-38.

Fertile pinnules of Recent Anemia vary considerably in habit. They may
be scarcely foliose, or broad and leaf-like. They may have flat segments, or
folded ones sometimes of boat-like appearance. The segments whether flat or
folded may be reflexed and the whole pinnule rolled backwards like a tightly clenched
fist, or they may be only slightly recurved or not reflexed at all. The species whose
pinnules compare most closely with the fossils in general appearance appears to be
the South American A. gardneri Hook., but in it the segments are fewer or more
tightly enrolled, less rigid and less conspicuously jagged at the margins than in
A. poolensis (PI. 32, figs. 11-13). The species A. schimperiana Presl. from Angola
also has closely folded pinnules in some herbarium material examined. A rare
species from Madagascar A. lanipes Christensen is somewhat illuminating. Norm-
ally, its pinnules appear to be flattened in one plane and they are so described by
Christensen. But a sheet No. 2798 (G. F. Scott Elliot Herbarium, Royal Gardens,
Kew, 986) shows some pinnules definitely reflexed and rolled in varying degrees.
In the most extreme instance the tightly rolled pinnules form sub-globular bodies
1*5 to 2 mm. in diameter. Perhaps this pinnule had become desiccated in life and
had curled as the result, thereby protecting the sporangia on its lower surface. A
similar tendency to protect by curling can be observed in British ferns growing in
situations liable to periodic desiccation such as walls or crevices.

In A. poolensis the rolling of the pinnules appears to be a more or less constant
character and it seems to be among terminal pinnules that rather flatter or less
recurved specimens are found. Presumably the rolled up " bodies " were torn
from the stipes by the battering they suffered from flood waters as they were swept
to the places of deposition among angular quartz sand and coarse woody fragments.

A . lanipes is a peculiarly hairy species. A . schimperiana is also covered by a close
felt of rather long hairs. The hairs of A. schimperiana are sparsely septate, long,
swollen at the septa with a transverse diameter of 20 to 30/4 across the cavity. Those
of A . lanipes and A . perrieriana Chr. are similar so that they differ definitely from the
fossil. Some Recent species have no hairs but are thickly beset with paraphyses
like those of the fossil, e.g. A. phyllitides and A. gardneri. Some species, like the
fossil, have both long hairs and turgid paraphyses, e.g. A . schimperiana. So far no
non-septate hairs have been found in the few living species examined. Often the

THE SCHIZAEACEAE OF THE SOUTH OF ENGLAND 303

paraphyses in the living are less convex on one side than the other, or they may be
concave on one side giving a curved form in profile. Others are symmetric. The
basal end is not infrequently more rounded and blunter than the distal, and attach-
ment is commonly by a short stalk either centrally placed or on one side. Paraphyses
in A. gardneri were no x 55/4 and in A. phyllitides 43 to 59/4 long, 1875 to 21-875/4
broad. A paraphysis wall was measured in A. schimperiana. In optical section it
was about 1-2/4 thick.

The spores of the fossil, A. poolensis, appear to be unique in having a smooth
glistening surface, but among the 115 living species of Anemia there are some whose
spores are still unknown, and others which it has not been possible to examine.
Nevertheless, many species were studied by Mr. Croft in the Herbarium of the
Botanical Department of the British Museum (Natural History). The majority show
highly characteristic ridged spores. The ridges may be smooth, tuberculate, or
spiny. In some species the spore outline is more or less triangular, in others
globose-tetrahedral. The size shows some little variation, e.g. 56 to 87-5/4 in A.
gardneri Hook, with a wall of about 4/4 thick ; 75 to 100/4 in A . schimperiana Presl. ;
9375 to 106-25/4 in A. adiantifolium Swz. ; 53 to 70-7/4 in A. perrieriana Chr., wall
3 to 5-8/4 thick ; about 81/4 in A. lanipes Chr. Thus they tend on the whole to be
larger than those of A. poolensis (50-60/4) which corresponds in size more closely
with those of the American species A. fremonti forma fertilis Andrews. These
according to Andrews & Pearsall (1941 : 169) show a considerable range from 25 to
47/4 with an average of about 40/4 ; some measured by the writer were as much as
50 or 53/4. The only smooth spores seen were immature examples of A . perrieriana.
They had thick firm walls. One plant gave smooth spores only, but the next on the
same sheet and no doubt from the same gathering had mature smooth-ridged spores.
In this species, as Mr. Croft noted, the spores become thick-walled prior to the forma-
tion of ornament. The ridges vary in their distance apart in different species and
also in their smoothness or degree of ornamentation as will be further discussed on
p. 306.

Thus while the apical annulus indicates the family Schizaeaceae beyond doubt,
the generic relationship calls for careful consideration. The reflexed enrolled fertile
pinnules, annulus with a single row of cells, many-celled apical plate, and spore
size, all point to Anemia as the nearest living genus, but the smooth unridged spores,
differing from the mature spores of known species, are peculiar to the fossil and require
some explanation. Can they possibly be regarded as immature spores like those of
A. perrieriana described above ? or as mature spores, originally ridged but denuded
in fossilization of an ornamental exine ? or should the fossils be separated on the
grounds of smooth spores and non-septate hairs as a distinct genus ? While there
can at present be no absolute proof of the generic status, the following considerations
appear to the writer to justify the use of the name Anemia at least provisionally.

(1) The sporangia have constantly (although not invariably) dehisced naturally
as if they were fully mature, but in undehisced specimens which have been protected
by the sporangium wall the spores show the same smoothness.

(2) Smoothness of spore is an invariable feature in every pinnule examined and is
seen not only in the Lake material but wherever the species has been found. It is

304 THE SCHIZAEACEAE OF THE SOUTH OF ENGLAND

most improbable that only immature material would have been found in different
localities and among so many different gatherings.

(3) Against the view that an ornamental exine has been lost in fossilization is the
fact that a second species from Colwell (see p. 305) has typical ridged spores although
this material has been far more macerated in fossilization than that now under
consideration.

(4) In two fossil species of Lygodium hereafter described (pp. 308, 312), one
has consistently smooth spores, the other has highly ornamental spores while the
majority of living species of Lygodium have spores ornamented in various distinctive
ways, though L. elmeri, for example, has smooth spores. It seems unreasonable,
when smooth and ornamental spores can both occur within a related genus, to make
smoothness a reason for separating A. poolensis generically from living Anemia.
Possibly smoothness of spore is a primitive character which still persists in Lygodium
to a small degree but is now lost in living Anemia so far as present knowledge goes.
On the whole the balance of probability appears to the writer to justify reference of
the fossil to the genus Anemia itself, although it must be admitted that the last word
may yet remain to be said.

Anemia is an herbaceous genus. Some examples of habitat are here quoted from
herbarium sheets at Kew: 1,500 metres in Angola, 3,000-4,000 ft. in Cuba, 1,500 ft.
in Mexico, 3,900 ft. in New Granada, 5,000 ft. in Madras in the Nilghiri Hills. There
are records of " shady rock crevices", "summit of dry mountain ranges", " wet
rocks", " by the river ", "on rich moist shady banks under shrubs ", " moist cal-
careous river bluffs ", crevices in vertical side of limestone rock " and " rocky pine-
woods ".

Anemia colwellensis n. sp.
(PI. 35, figs. 39, 40, 42-53 ; PI. 36, figs. 59-64 ; Text-fig. 2, i)

DIAGNOSIS. Fertile pinnule flat, about 2-5 mm. long, 17 mm. broad, with several
(at least 8 ?) rounded alternating segments on opposite sides of a nerve. Free outer
margins of segments sharply recurved on to the lower surface so as to give a smooth
outline and to form a false fringed indusium. Main vein of pinnule sinuous.
Nervation of segments flabellate, the nervules repeatedly forked and free. Upper
surface smooth, without hairs ? Sporangia in two rows on the lower surface of each
segment, sub-ovoid, somewhat laterally compressed giving a slight bisymmetry ;
sessile, broadly attached by the whole breadth of the base. Length of sporangium
very variable, maximum about 228 to 342;* ; breadth about 171;*. Annulus slightly
bisymmetric, apical, conspicuous, formed by a single row of about 15 cells with
extremely thick walls, occupying about one-half to one-third of the length of the
sporangium. Trilete tetrahedral-globose spores ornamented with ridges which
give the effect of tubercles where seen in profile at the margins. Diameter of spores
37-5 to 66-6/4, commonly 43 to 50/4. Height of spores from back to front 37-5 and
43 '75/* m two specimens measured. Distance between ridges about 3 to 6-25^.
Paraphyses many among the sporangia, sessile or shortly stalked, about 114 to I2O/*
long, 48 to 72;* broad.

HOLOTYPE. ¥.31519-23.

THE SCHIZAEACEAE OF THE SOUTH OF ENGLAND 305

DESCRIPTION. Fertile pinnule 2*5 mm. long, 1-7 mm. broad, having several (at
least eight) short, rounded, closely contiguous segments situated alternately on the
opposite sides of a nerve and inclined to it at angles of about 45° (Text-fig. 2, i).
The free outer margins of the segments are sharply recurved on to the lower surface
thereby giving the peculiarly smooth, rounded outline shown and producing a flat
false indusium on the lower surface, which partially covers the sporangia. The
" indusium " is striate and marginally fringed, the alignment of the fringe being
parallel with the planes of separation between the segments and corresponding with
the cell-structure (Text-fig. 2, i). Main vein of pinnule sunk in a sinuous channel on
the upper surface, not actually seen on the lower where it was obscured by sporangia
and masses of released spores. Nervation of segments flabellate, the venules being
repeatedly forked and free as in the living Anemia and as in Anemia sp. 3 (see PI. 36,
fig. 65). Upper surface smooth and free from hairs, lower surface with turgid
brown sub-fusiform paraphyses among the sporangia showing, when highly magnified,
a curious transverse streaking perhaps due to stretching caused by crushing as it is
not always apparent. The paraphyses may be shortly stalked but are usually rather
broadly based. They are thin-walled and about 114 to 120/1 long, 48 to 72 p broad
(PI. 35, figs. 47, 48). One fragment of pinnule segment showing the distal margin
appeared to be at least two cells thick and displayed oblong, almost rectangular,
cells about 6 to 12/4 in diameter with very fine closely digitate (sinuous) walls. The
free ends of the nervules in this fragment were about 12/4 apart.

The sporangia (PL 35, figs. 39, 40, 42-46) are large although they vary considerably
in size with the degree of development and possibly with their position on the pinnule.
The largest are about 228 to 342/4 long and about 171/4 broad. They are sub-ovoid
but somewhat flattened so as to be bisymmetric, sessile and broadly attached by
the whole breadth of the base, tearing away from the surface of the pinnule in a
ragged manner often with fragments of epidermis still adhering. The apical annulus
is in some cases somewhat bisymmetric owing to the flattening, it is a conspicuous
feature sometimes bulging out beyond the sporangial wall below, occupying from
one-third to one-half the length of the sporangium. It consists of a single row of
at least 15 long narrow cells with straight thick walls which fork conspicuously at
their lower extremities where they alternate with the thinner cells of the sporangial
wall. Dehiscence longitudinal with stomium long and conspicuous. Its cell-
structure is obscure owing to the denseness of the material and its broken character.
It appears to have been directed towards the margin of the segment when in the
position of growth. Apical plate multicellular, delicate, of angular equiaxial cells.
Cells of sporangial wall elongate parallel to the major axis of the sporangium (PI. 35,
fig. 45), about 24/4 broad, straight superficially, unevenly thickened so as to have
a " beaded " appearance when seen from the interior. They contain numerous
rounded or angular bodies which may have been chloroplasts but they are now heavily
pyritized and glistening (PL 35, figs. 42, 43, 45). Spores (PL 35, figs, 49, 53 ; PL 36,
figs. 59-64) trilete, tetrahedral-globose, somewhat triangular in outline, conspicu-
ously but finely ridged, the ridges when seen in profile giving a false tubercled appear-
ance to the outline of the spores. Distance between the ridges varies in different
parts of the spore and may be from 3 to 6-25/4. The distal ends of the rays sometimes

GEOL. II, 7. 25

306 THE SCHIZAEACEAE OF THE SOUTH OF ENGLAND

fork, and splitting may occur along the rays (PI. 36, fig. 62). The diameter of the
spores varies from 37-5 to 66 -6ju, and is commonly from 43 to 50;*. The height from
the dorsal to the ventral side is more or less equal to the diameter. Two measure-
ments of height were 37-5 and 43 75/4.

REMARKS. One fertile pinnule more or less complete. This minute fragment
was so tender that it fell to pieces on attempting to lift it on to a slide. Fortunately
a drawing of the under surface had been made and a preliminary examination of
both surfaces. Even more fortunately it was possible to make several slides of the
remains showing sporangia, annuli and spores. Paraphyses are also preserved on
the slides, but there are no indications of a felt of hairs. A small fragment of the
pinnule itself is mounted on one slide showing nerves and cell-structure. The slides
are rather opaque as the material was too scarce to risk destroying it by more than a
slight degree of maceration in nitric acid. The character of pinnule, sporangia and
and spores indicates the genus Anemia without question. Fortunately, in this
case, the spore ornamentation is typical of Anemia. It is significant that the orna-
mentation is preserved in spite of the rotten and fragile state of the pinnule fragment,
so much more rotted in fossilization than the pinnules of A. poolensis. The flat
unreflexed pinnule with fringed false indusium formed by the reflexed margins of the
short smooth-edged segments, the short broadly attached sporangia without hint
of narrowing or arising from a stalk, the large deep annulus and the ridged spores,
distinguish this species without any doubt at all from A. poolensis. Study of some
of the many living species of Anemia indicates that the ridges on the spores vary
to some extent in different species in their distance from one another and in their
ornamentation. Thus in A. schimperiana the ridges are simple, not tubercled, and the
spores sharply triangular with clear translucent rounded projections at the angles.
The ridges are about 6/1 apart. In A. gardneri the ridges bear contiguous tubercles
which are a conspicuous feature, the tubercles being about 6/t in diameter. In
A. adiantifolium the spore has round translucent angular projections, the ridges are
smooth, and about io/* apart. In A. lanipes they are sharp and thin, and may be
6/* apart, although sometimes closer. A. aurita Sw. has ridges with tubercles, A.
anthriscifolia Schrad. has smooth or minutely echinulate ridges. In A. perrieriana
the surface of the spore is ridged coarsely. A. affinis Bak. has large spiny spores.
The ridges on the fossil A. fremonti forma fertilis are about 2-1 to 2-5/4 apart.
A . phyllitides has spines or long pointed tubercles arranged on ridges in the mature
spores. It would involve prolonged study to attempt any sort of classification of
the genus by its spore characters or even to prove conclusively that such was possible.
Such a study cannot be undertaken for the purposes of this paper but knowledge of
Anemia is incomplete without it.

The pinnule segments of the fossil recall A. imbricata Sturm. (Brazil), A.ferruginea
— a hairy species from Central America (= A. fulva Hook.), the flat pinnules of
A. lanipes (Madagascar), and A. tomentosa Sw. (warm America from Mexico and the
Antilles to Peru and the Argentine). More fossil material is needed to complete
the record of this interesting fern. But whatever view may be taken of the generic
position of A. poolensis. the presence of Anemia in Southern England is indisputably
established by the tiny fertile fragment described above as A. colwettensis.

THE SCHIZAEACEAE OF THE SOUTH OF ENGLAND 307

Anemia sp. 3
(PI. 36, fig. 65)

1926. Filix incertae sedis, sp. 5 ; Reid & Chandler, p. 39, pi. i, figs. 10, n.

To the description previously published may be added the additional fact that
some of the pinnules are asymmetric. There can be little doubt that this fern belongs
to Anemia. For details of pinnules and nervation see Reid & Chandler (1926). The
characters of these barren pinnules closely resemble those of the pinnule segments of
A. colwellensis both in the smoothness of their margins and in the flabellate nervation.
It is interesting to speculate whether the Bembridge specimens can possibly be barren
material of A. colwellensis, but there the matter must rest until further evidence is
available to prove or disprove their specific identity.

Genus LYGODIUM Swartz

Lygodium prestwichii (Gardner & Ettingshausen)
(PL 36, figs. 66-69)

1854. Asplenium sp.? Prestwich, p. 156, pi. 3, fig. 6.

1880. Pteris (?) prestwichii Gardner & Ettingshausen, p. 53, pi. 10, fig. 8.

1886. Lygodium prestwichii (Gard. & Ett.) Gardner, pp. 400, 401, 403, pi. 2, figs. 2-4.

DIAGNOSIS. Pinnae simple, elongate lanceolate-linear or cleft. Veins diverging
at a sharp angle from the midrib, crowded, forking once or twice, midrib considerably
thinned towards the apex. Fertile pinnules borne around the toothed margins of
the barren pinnules ? Sporangia, spores and fertile pinnules unknown.

NEOTYPE. ¥.24862.

REMARKS. Fragments of this fern were found at Counter Hill, Lewisham, by
Prestwich (1854, pi- 3> fig- 6 ; see also Gardner & Ettingshausen 1880, pi. 10, fig. 8),
and by Gardner at Croydon and Woolwich (Gardner 1886, pi. 2, figs. 2-4). Gardner
considered all these specimens as identical specifically after comparing his own
material with that of Prestwich (1886 : 403). He regarded the species with its
sparsely-toothed pinnules as characteristic of the Woolwich Beds and as quite
distinct from Lygodium kaulfussi from Bournemouth. One fragment figured by
Gardner (1886, pi. 2, fig. 2) shows a " toothed " margin, the projections being (so he
states) the bases of fertile segments removed hi fossilization. Unfortunately the
Prestwich pinnule from Lewisham seems to have disappeared (Gardner & Ettings-
hausen, 1880 : 53) and even Gardner's Woolwich and Croydon specimens cannot
now be traced. It is therefore fortunate that a few fragmentary pinnules with
toothed margins which apparently belong to the same species were collected more
recently by the late H. C. Berdinner in beds at Newington. From the locality and
their appearance they may be presumed to be of Woolwich age. They are impres-
sions only representing fragmentary pinnules. One (¥.24862) with counterpart,
shows two lobes of a pinna, one lobe being much puckered and folded longitudinally.
The midrib is stout and tapering, rounded and prominent on the lower surface

308 THE SCHIZAEACEAE OF THE SOUTH OF ENGLAND

(represented by a furrow on the impression). The nerves are about 0-6 to 0-75 mm.
apart where they spring from the midrib. They are represented on the impressions
by thin raised thread-like lines. The marginal teeth are well preserved in parts
(PL 36, figs. 66-68) ; the margins of the frond were thickened, especially at the tips
of the teeth. There is nothing in this specimen, or in ¥.24861 which also shows
teeth, to indicate that the teeth were associated with fallen fertile pinnules as Gardner
supposed, and on the whole the appearance of the two specimens is against this
interpretation. Nevertheless it is possible that these particular fragments happened
to have been barren, whereas if they had been fertile the association of teeth and
fertile pinnules might have been shown. But whether the teeth were, or were not,
associated with fertile pinnules, it seems clear at least that their presence indicates
that in the Woolwich Beds there was a species quite distinct from the Bagshot
L. kaulfussi with its entire-margined barren pinnules and its much-branched fertile
pinnules which apparently were borne in a panicle at the tips of barren fronds and
not around their margins. In describing the species Gardner (1886 : 403) recorded
that " the pinna was simple or cleft into two or more lobes". The veins were free,
diverging at a sharp angle from the midrib, forking once or twice. Prestwich
figures nerves about 0-75 mm. apart near their origin on the midrib.

Gardner thought the species indistinguishable from Lygodium japonicum Sw. a
native of Japan, China, Ceylon, Java and the Philippines, but this is a curious con-
clusion for it does not accord with his views about the position of the fertile pinnules
of the fossil. In living L. japonicum they are borne in clusters at the tips of the barren
pinnules as they are also in L. articulatum Rich, and L. palmatum Sw. L. prestwichii
rather resembles the other group of living species in which the strap-shaped pinnules
whether simple or parts of subdivided pinnae carry the fertile pinnules around
their margins, an arrangement seen, for example, in the living L. flexuosum (L.),
L. volubile Sw., L. circinnatum (Burm.) and L. salicifolium Presl.

Lygodium is a climbing fern of tropical and subtropical lands in both hemispheres.
In North America it occurs in Florida and Massachusetts in woodlands and in low-
land areas of New Jersey. In South America it extends to southern Brazil east of the
Andes but less far south on the west of the mountain range. Professor Halle (1940)
points out that his fossil species L. skottsbergii is found 30° south of the present
extension of the genus. In the old world the genus goes north into China, e.g.
500 metres above sea level in Szechuan and southward to New Zealand, occurring
throughout the whole of the tropical belt. Some species have pinnules with toothed
margins and in others the margin is entire.

Lygodium kaulfussi Heer
(PI. 37, figs. 70-83 ; PI. 38, figs. 84-87)

1861. ? Lygodium kaulfussi Heer, p. 409, pi. 8, fig. 21 ; pi. 9, fig. i.

? 1879. Asplenites prae-allosuroides Gardner & Ettingshausen, p. 34, pi. 3, figs, i, 2.

1880. Lygodium kaulfussi Heer : Gardner & Ettingshausen, p. 47, pi. 7, figs, i, 3-8 ; pi. 10,

fig. ii.

1882. Lygodium kaulfussi Heer : Gardner, pp. 67, 68.
1886. Lygodium sp., Gardner, pp. 401, 404, pi. 3, fig. 9.

THE SCHIZAEACEAE OF THE SOUTH OF ENGLAND 309

DIAGNOSIS. Barren fronds simple or deeply palmately divided, tapering to the
base. Fertile pinnules borne in a sympodially branched tuft with segments in
pairs or threes. Spores trilete and tetrahedral-globose, shining, smooth or minutely
granular, about 75 to 112/4 in diameter.

NEOTYPE. ¥.14962.

DESCRIPTION. Barren pinnules have been described and figured from Bourne-
mouth and Studland, and two small tufts of fertile pinnules from Bournemouth
(Gardner & Ettingshausen, 1879, 1880 ; Gardner, 1886). Gardner states that the
barren pinnules from Studland were very near to those from Bournemouth but with
" much closer veins", and suggested that the Studland specimens were intermediate
in form between L. kaulfussi from Bournemouth and L. prestwichii from the Wool-
wich Beds. It is not clear from his statement whether he really regarded the Studland
material as belonging to a species distinct from L. kaulfussi. But fertile pinnae
with spores now afford evidence that the Studland and Bournemouth material is
specifically identical (cf. PI. 37, fig. 78 ; PL 38, fig. 86 from Branksome Dene, with
PI. 37, figs. 72, 74, 80-83; PI. 38, fig. 85 from Studland). Moreover, the recently
collected barren pinnules from Studland agree closely with some of Gardner's Bourne-
mouth specimens. The differences of nervation which he noted are therefore
probably individual variations with no specific significance. This view is supported
by the variations which can be found in living Lygodium species.

Gardner & Ettingshausen' s description of the Bournemouth fronds is full and
detailed, and it is not necessary to repeat it, but Gardner gave only the briefest
account of the single Studland fragment figured. The following additional details
based on the fresh material may therefore not be out of place.

The barren fronds are not infrequently to be found as perishable impressions in
the lower part of the soft laminated dark grey clays about 4 to 5 ft. above the sand-
rock, i.e. within the lowest 100 ft. of the Lower Bagshot Beds. Often the impres-
sions are covered by decayed remains of actual fronds which quickly crack and
crumble on exposure to air. The few pinnules so far seen are variable in shape
and sometimes divided near the base (PI. 37, fig. 70). The lobes may be short or
long and lanceolate, obtuse at the apex, tapering somewhat towards the base, with
entire margins. The primary nerves are stout but become attenuated above, the
secondary nerves are close together, about 0-4 to 0-5 mm. apart, given off at acute
angles, thrice forked, sharply defined.

Fertile pinnules (PI. 37, figs. 72-78). The specimen figured from Bournemouth
(Gardner & Ettingshausen, 1880, pi. 10, fig. n) shows a repeatedly divided pinna
with stalked pinnules which are linear-elongate and serrate along the margins.
Gardner & Ettingshausen (1880 : 48) state that the pinnules in the upper part were
" solitary or undivided, but in the lower part two or three occur together or are twice
or thrice divided ". This two- or three-fold division may also be seen in the living
L. articulatum Rich. The midribs are sinuous and the rachides unwinged. Gardner
did not record sporangia nor spores, but a preparation from ¥.14962 (the neotype)
has revealed the presence of spores identical with those described below.

In 1882 (p. 68) Gardner reconsidered the specimen which he and Ettingshausen
(1879) na<3 named Asplenites prae-allosuroides (¥.14968), and decided that it was a

3io THE SCHIZAEACEAE OF THE SOUTH OF ENGLAND

fertile frond of Lygodium with " exceptionally abbreviated pinnules". No spores
could be obtained from this specimen when it was re-examined recently, and in
view of the difference in form of the fertile segments its identity with L. kaulfussi
must remain doubtful.

The fertile pinnules from Studland are well preserved. They were isolated from the
matrix by boiling, washing and sifting, and were then examined microscopically.
Only detached solitary fertile pinnules have as yet been seen. When perfect they
are petiolate and, as in the Bournemouth specimens, without any wing-like extension
of the lamina along the stalk. They are linear-elongate, obtuse at the apex, serrate
along the margins, while the markedly sinuous midrib is sometimes very conspicuous,
although only slightly raised, on the upper surface; it is often prominent on the lower
surface which may have a longitudinal facet on each side of it (PI. 37, figs. 73, 75).
The epidermis is formed of deeply interlocking cells with beaded appearance due to
unevenly thickened walls (PI. 37, fig. 79). The diameter of the largest cell seen
including the digitations was 0-048 mm. The sporangia are oval, flattened, sunk
in oval hollows on the underside of the pinnules, concealed by overlapping bracts
or indusia. They remain obscure even in macerated specimens because of the thick
and opaque character of the bracts and pinnules which surround and protect them.
They diverge in a pinnate manner from the midrib, one sporangium corresponding
to each tooth of the serrate margin. The walls of the sporangia are usually ruptured
so that as a rule the form, structure and annulus are destroyed or masked, especially
if covered and surrounded by a mass of liberated spores. After maceration, remains
of the annulus can sometimes be seen with thick-walled more or less parallel-sided
elongate cells.

The spores (PI. 37, figs. 80-83 ; PI- 3$, figs. 84-87) are numerous, globular-tetra-
hedral with clear fine triradiate marks, the rays sometimes forking slightly at their
free extremities. The surface is shining and smooth or very finely granular resembling
figures of L. skottsbergii Halle (1940, pi. i, figs. 9, 10) from supposed Eocene beds of
Coronal, province of Concepcion, South Chile, South America. A few tubercles or
granules are sufficiently prominent to be visible in profile, but they are probably
fragments accidentally adhering to the surface, for although apparent when the
spores are first mounted in glycerine, they quickly disappear leaving a perfectly
smooth surface. The diameter of the spores varies from 75 to 1127*, and is frequently
about ioo/i when uncollapsed, very rarely larger. The thickness of the coat in
the largest spore seen was as much as 6/t. More frequently it is only about 3/4 in
optical section. The length of the fertile pinnules is about 4-5 mm. ; breadth about
1-5 mm. Length of toothed segments of fertile pinnules about 0-5 mm. Identical
fertile pinnules with identical spores were also found at Branksome Dene, Bourne-
mouth Freshwater Beds (PI. 37, fig. 78 ; PI. 38, fig. 86).

It must be stressed that barren and fertile material have not yet been found in
organic connexion but as their association in the same beds at Studland almost
certainly points to derivation from a single species, they are so regarded and described
here. The fertile pinnules and spores afford conclusive evidence that the Studland
and Bournemouth material is specifically identical, while the barren fronds from the
two localities appear to the writer to agree also,

THE SCHIZAEACEAE OF THE SOUTH OF ENGLAND 311

An isolated spore, apparently of this species, has also been found in the Cliff
End Beds of Mudeford (Auversian ?) (PL 38, fig. 87). Halle (1940: 261) pointed
out that the type of L. kaulfussi Heer was an indeterminable and useless fragment
whereas the numerous fossil records of L. kaulfussi are, in fact, based on the well-
preserved English Eocene fronds unfortunately identified with the useless Skopau
type. Hence, he suggested, Gardner's specimens may, for practical purposes,
serve as the type of the species L. kaulfussi and the specimen British Museum
(Natural History) No. ¥.14962 is here designated as the Neotype.

The Studland and Bournemouth fossils have been united as a single species only
after careful consideration of the evidence afforded by the fertile pinnules from both
localities following upon a study of the spores of twenty-six living species. This
investigation was made possible by the kindness of Mr. A. H. G. Alston of the
Botanical Department, British Museum (Natural History) . It strongly suggests that
when their characters have been fully studied they will prove to be sufficiently
distinctive for use in specific determination in the majority of species. But much time
and labour would be required in order to work out the spore characters of the genus
in a really satisfactory manner. For in every species it would be essential to examine
a wide range of material from different plants and localities to find out the true
limits of variation of the spores within each. Moreover the determination of the
Herbarium material itself would have to be checked. As an illustration of the nature
of the problem it is perhaps sufficient to quote the cases of L. volubile Sw. or L.
circinnatum (Burm.) which displayed considerable variations of spore character on
different sheets and which perhaps include more than one species in each case. On
the other hand readily recognizable species such as L. reticulatum Schk., L. articu-
latum, L. japonicum, L. scandens (L.), L. polymorphum (Car.), L. smithianum Pr.
and L. palmatum Sw., each have distinctive spores which appear constant in character.
The spores of different species may vary in form, e.g. they may be more or less conical,
or sub-globular, or more or less sharply angled. They also vary in size from about
40/4 in diameter in L. borneense and L. elmeri ; 40 to 50/4 in L. japonicum and L.
polymorphum ; and 80 to over ioo/* in L. hians, L. articulatum, L. reticulatum, L.
cubense, L. merrilli and L, lucens.

Whereas by far the greater number of living species have spores which are orna-
mented in a greater or lesser degree they do vary considerably in surface ornamenta-
tion and thickness of spore wall, while a few species are smooth and shining. Thus
L. elmeri is smooth and fairly thick-walled with only very occasional scattered minute
tubercles. L. polymorphum is thin-walled and finely granular all over. L. reticu-
latum has a coarse reticulation of the surface, and L. scandens is somewhat similar
with a series of deep pits. In L. articulatum the whole surface is covered by low,
rounded, contiguous tubercles which diminish in size between the arms of the three-
rayed scar on the ventral surface. L. japonicum is evenly covered with regular, low,
rounded, contiguous tubercles smaller than those of L. articulatum. L. borneense
is thin-walled with small distinct close-set tubercles. L. merrilli has larger but quite
distinctly separated tubercles. In L. smithianum and in one sheet attributed to
L. volubile (Nardil, 1891) from Trinidad, the spore has a fairly broad equatorial
flange, irregular scattered large wart-like tubercles sometimes as much as io/* broad,

312 THE SCHIZAEACEAE OF THE SOUTH OF ENGLAND

with finer tubercles between, especially on the surface in the angles of the rays.

The spores of L. kaulfussi from Studland and Bournemouth are characterized by
their large size, often 100/4 or larger, thin walls, and smooth or very finely granular
surface. These characters combined, together with the globose-tetrahedral shape,
appear to distinguish them from any living Lygodium seen. As already mentioned
(p. 310) smooth-walled spores of exactly similar appearance and comparable size
(65-105/4 commonly 85/4) occur in Halle's species L. skottsbergii. The fertile fronds
of this species appear to have had a mode of growth in a tuft like that of L. kaulfussi
but show a much reduced lamina on the sides of the rachides, absent (possibly
through lack of preservation) in L. kaulfussi. Halle separated his species from L.
kaulfussi on account of the shorter, broader segments of the barren pinnules, their
asymmetrical shape and truncate base, and especially on account of the number of
segments (three). But he writes that the fertile specimens of L. skottsbergii might
belong to L. kaulfussi. There is certainly a close general resemblance between the
two, and this is supported by the character of the unusually smooth spores which
are also comparable in size and shape. They are certainly more similar to one another
than to any living species, and it is tempting to think that they may be identical
and that such variations as occur may even be the result of different climatic or
ecological conditions.

A few smooth spores and barren and fertile pinnules attributed to L. gaudini
Heer are figured by Gilkinet (1922 : 5, pi. 14, figs. 2-9). Unfortunately the figures
show little detail and spore dimensions are not given. The material is from beds
of uncertain age in the argiles of Andenne (Aquitanian or Lower Miocene ?) yielding
also a number of Bovey Tracey species. There is no obvious difference from L.
kaulfussi although Gardner regarded Heer's two species (L. kaulfussi and L. gaudini)
as quite distinct. Possibly in view of Halle's comment on the first named (see p. 311)
they should not be separated. It is tempting to wonder whether in view of the pre-
ceding remarks L. kaulfussi may have been a plant of very wide range, particularly
in Eocene times, possibly even a persistent relic of the ancient more or less world-
wide genera of the tropical and subtropical province.

While, however, the specific relationship of these forms from wide-spread localities
must await further confirmation from an exhaustive comparative study of the actual
fossil material, there can be no reasonable objection to uniting the Studland and
Bournemouth fossils of which such a study has been made.

The general distribution of Lygodium is given on p. 308.

Lygodium poolensis n. sp.
(PI. 38, figs. 88-96)

DIAGNOSIS (based on spores). Spores trilete, subglobular-tetrahedral, ranging
from about 50 to 70/4 in diameter, commonly about 65 to 68/4. Depth from the
ventral to the dorsal side about 41-42/4. Surface ornamented with numerous flat-

THE SCHIZAEACEAE OF THE SOUTH OF ENGLAND 313

topped tubercles about 4/1 in diameter below, projecting conspicuously from the
surface for about 4/4 as seen in optical section.

HOLOTYPE. V.3I533.

DESCRIPTION. Barren pinnules unknown.

Fertile pinnules known only from a single naturally macerated fragment about
175 mm. long and from a detached spore on a slide of Anemia. Maceration has
removed the lamina leaving the slightly sinuous midrib and a few attached solitary
sporangia which are borne on it in a pinnate manner at the ends of the nervules.
One sporangium (PI. 38, figs. 88, 89) is perfect, four others are incomplete, two of
them being almost entirely broken away. The perfect sporangium is broadly
bolster-shaped or sub-ellipsoid with a large spreading annulus on one side near the
proximal end. It was very clearly displayed when the specimen was first placed in
nitric acid and can still be distinguished when examined by strong reflected light
against a white background. The sporangium is about 0-55 mm. long, 0-3 mm.
broad. The annulus about 0-34 mm. in maximum diameter extending for over half
the length of the sporangium. It is only about 0-17 mm. in depth.

Spores numerous, trilete, tetrahedral-subglobose, low conical or low triangular-
pyramidal having a sub-circular or rounded triangular outline. Triradiate mark
clear, sharp and fine, the rays about 18-25/4 in length. Surface highly ornamental
having a series of deep pits whose coalescent walls produce the effect of rather flat-
topped tubercles (PI. 38, figs. 90-96). Diameter of tubercles often about 4^ ; height
2-5 or 3 to 4/4. About four tubercles occupy a square of about 12-5 x 12-5/4. The
tubercles project conspicuously at the circumference of the spores when seen in
profile. Typical spore dimensions are 65/4 ; 70 x 50/4 ; 65 x 60/4 ; 55 x 687/4 ;
66 x 50/4 ; 66 x 57/4 ; 57 x 54/4 ; 63 x 48/4. One spore which measured 687/4
in diameter was 41-25/4 in height (dorsi- ventral profile) ; thickness of spore walls
where measured about 4/4.

REMARKS : As already noted there are some grounds for regarding spores of
Lygodium as of use in specific determination. Tubercled spores similar in character
to those of the fossil occur in very few living species but are present in L. scandens
Sw. and L. reticulatum Schk. These two species are grouped together by Diels on
account of their reticulated spores which appear to separate them from other species.
A sheet of L. scandens (Thwaites 1404) in the Herbarium of the Botanical Depart-
ment, British Museum (Natural History) showed spores varying in diameter from
about 66 to 84/4, and in another which more closely resembled the fossil (Ross 123
from Nigeria) spore diameter was 60 to 84/4 and the height about 41/4 (PI. 38, figs.
97-101). The tubercles appeared somewhat less conspicuous and prominent at the
margin of the spore as well as rather coarser and more rounded than in the fossil.
A sheet of L. reticulatum (Compton 244) had larger spores, 80 to no/* in diameter,
while another sheet from New Hebrides (McGillivray 1860) showed similar variations
of size. The tubercles were considerably less prominent at the margin than in the
fossil.

Diels gives the geographical range of L. scandens as West Africa, Further India
to Chitral, South China, Malaya to North Australia, and of L. reticulatum as North
Australia, Melanesia and Polynesia. Hence the fossil L. poolensis appears to be

GEOL. II, 7. 16

314 THE SCHIZAEACEAE OF THE SOUTH OF ENGLAND

related to an essentially tropical old world sub-division of Lygodium. In L. scandens
the nerves of the barren pinnules are free ; in L. reticulatum they anastomose. A
note on the sheet of L. scandens, from which spores are illustrated, states that it
occurred in swamp forest near sea level in Nigeria. L. poolensis is readily distin-
guished by its spores from the smooth-spored L. kaulfussi.

REFERENCES

ANDREWS, H. N. 1941. On the Flora of the Frontier Formation of Southwestern Wyoming.

Ann. Mo. bot. Gdn, St. Louis, 28 : 165-192, pis. 1-7.
CHRISTENSEN, C. 1932. The Pteridophyta of Madagascar. Dansk. bot. Ark., Kjobenhavn, 7 :

xv + 253 pp., 80 pis.
DIELS, L. 1902. Schizaeaceae. In Engler, H. G. A. & Prantl, K. A. E. Die Naturlichen

Pflanzenfamilien, 1, 4 : 356-372. Leipzig.

GARDNER, J. S. 1886. Fossil Plants of the Tertiary and Secondary Beds of the United King-
dom. Rep. Brit. Ass., Aberdeen 1885, : 396-405, pis. 1-3.
& ETTINGSHAUSEN, C. 1879. A Monograph of the British Eocene Flora, I . Filices : 1-38,

pis. 1-5. Palaeontogr. Soc., London.

1880. Ibid., II. Filices : 39-58, pis. 6-n. Palaeontogr. Soc., London.

1882. Ibid., III. Filices : 59-86, pis. 12, 13. Palaeontogr. Soc., London.

GILKINET, A. 1922. Plantes fossiles de 1'argile plastique d'Andenne. Mem. Soc. geol. Belg.,

Liege, 4 : 23-38, pis. 14-17.
HALLE, T. G. 1940. A Fossil Fertile Lygodium from the Tertiary of South Chile. Svensk

bot. Tidskr., Stockholm, 34 : 257-264, pi. i.
HEER, O. 1861. Beitrage zur nahern Kenntniss der Sachsisch-thiiringischen Braunkohlen-

flora. Abh. naturw. Ver. Sachs. Thiir., Berlin, 2 : 403-428, pis. i-io.
LANG, W. H. 1929. On Fossil Wood (Dadoxylon hendriksi, n. sp.) and other Plant-remains

from the Clay-Slates of South Cornwall. Ann. Bot., London, 43 : 663-681, pis. 15, 16.
PRESTWICH, J. 1854. On the Structure of the Strata between the London Clay and the

Chalk in the London and Hampshire Tertiary Systems. Part II. The Woolwich and

Reading Series. Quart. J. Geol. Soc. London, 10 : 75-170, pis. 1-4.
REID, E. M. & CHANDLER, M. E. J. 1926. The Bembridge Flora. Catalogue of Cainozoic

Plants in the Department of Geology, I. viii + 206 pp., 12 pis. Brit. Mus. (Nat. Hist.),

London.
SAPORTA, G. de. 1868. Prodrome d'une flore fossile des travertins anciens de Sezanne.

Mem. Soc. geol. Fr., Paris (2) 8 : 289-436, pis. 22-36.

1 OCT 1955

PLATE 32

Anemia poolensis n. sp.

FIG. i. A recurved pinnule with attachment to stipe at (a). Several segments are seen
directed downwards with the sporangia on the lower surface projecting along their margins.
The transverse and longitudinal alignment of the cells over the middle of the segments on the
upper surface is clearly seen, x 26. ¥.31495.

FIG. 2. A single boat-shaped segment of a pinnule, side view with upper surface on the right.
It shows the crowded sporangia arranged in two rows on the under surface, x 15 (now dis-
integrated) .

FIG. 3. The same, enlarged. The apical annuli with elongate cells can be distinguished.
X ca. 35.

FIG. 4. A segment, lower surface with distal end at the top of the figure. It shows the
sporangia springing from the pinnule close to the nervule. Annuli of sporangia and elongate
cells of sporangial wall and pinnule are visible, x 50. ¥.31496.

FIG. 5. Segment attached at (a), dorsi-lateral view showing the apices of three sporangia
which project beyond the margin of the segment. Two have gaped longitudinally. The
sporangium at the distal end (top in figure) is shown in PI. 33, fig. 15. x 28-5. V. 31497.

FIG. 6. A pinnule segment covered with a felt of pointed hairs. Attachment at (a), x 50.
¥.31498.

FIG. 7. A detached sporangium showing the contraction to the attachment and short stalk
at (a). The dense region of the annulus is visible at the opposite extremity, x 150. ¥.31499.

FIG. 8. Another with dark thick-celled annulus and attachment (twisted) at (a). The thin
cells of the sporangial wall show the bead-like thickening which is present on the inner surface
X 150. ¥.31500.

All the above are from Lake, Dorset.

FIG. 9. Apical view of a disrupted annulus with stomium at (s). Remains of the torn
sporangial wall close to the line of the stomium are seen at (spg) . x 150. ¥.31516. Sandbanks,
Dorset.

Fig. 10. Apex of a sporangium, side view, showing the thick-walled annulus and some of the
contained spores. Granules in the cells of the sporangial wall chiefly below the annulus appear
to be chloroplasts. x 150. ¥.31512. Branksome Dene, Dorset.

Anemia gardneri (Recent)

FIG. ii. Part of a pinna to show the habit, with tertiary reflexed pinnae arising from the
stipe and bearing reflexed enrolled pinnules, x 5-5.
FIG. 12. The same, opposite side, x 5-5.
FIG. 13. The same (upper part) to show the recurved pinnules more clearly, x 15.

Bull. B.M. (N.H.) Geol. 2, 7

PLATE 32

10

ANEMIA POOLENSIS A. GARDNERI (Recent)

PLATE 33

Anemia poolensis n. sp.

FIG. 14. Apex of a dehisced sporangium showing the stomium and the thick-walled annulus.
A few cells of the broken apical plate still adhere to the upper ends of the annulus cells. A few
cells of the sporangium wall with bead-like thickening are seen at sw. x 150. ¥.31501.

FIG. 15. Apex of a sporangium which has dehisced longitudinally. It shows clearly the ring
of thick-walled cells which form the annulus and the thin-walled cells bordering the stomium
(st). The multicellular apical plate lies between the upper ends of the annulus cells. Spores
still remaining within the sporangium are visible at s, s,s. x 161 . This sporangium is the top-
most one shown in PI. 32, fig. 5. ¥.31497.

Fig. 1 6. The apex (side view) of a sporangium (sp i) of which the apical plate is obscurely
seen at (a). (an) is the thickened annulus. Part of the annulus of another sporangium with its
apex to the right is at (sp2). x 150. ¥.31502.

FIG. 17. The sporangium (spi) in fig. 16 showing the cells of the burst apical plate (a)
between the upper ends of the thickened annulus cells, x 400.

FIG. 1 8. Part of a pinnule segment. Two paraphyses (p, p) are still in position on the under
surface which faces the camera, x 75. ¥.31503.

FIG. 19. A detached almost unmacerated paraphysis showing the inflated form and opaque
character before maceration, x 400. ¥.31504.

FIG. 20. Another large detached paraphysis which has begun to collapse and crumple as
the result of maceration. The opaque lining has begun to break down as can well be seen at the
proximal end (p). The point of attachment was at (a), x 400. ¥.31499.

FIG. 21. Another (still opaque) showing the point of attachment below at (a), x 400.

V.3I505-

FIG. 22. Another after treatment, translucent but still inflated although somewhat crumpled.
X 400. ¥.31506.

Anemia schimperiana (Recent)
FIG. 23. A Recent paraphysis for comparison (Gossweiler 9705). x 400.

Anemia poolensis n. sp.
FIG. 24. A spore showing the smooth inflated form and (obscurely) the three rays, x 400.

FIG. 25. Another, partly out of focus, showing two of the three rays clearly. The smooth
inflated form is again shown, x 400. ¥.31505.

FIG. 26. The same spore with different focus. The thickness of the wall and smooth rounded
form are indicated. The rays are out of focus, x 400.

FIG. 27. Another spore showing the triradiate mark. The forking of the free ends of the
rays is clearly visible, x 400. ¥.31508.

All the above, except fig. 23, are from Lake, Dorset.

FIG. 28. A spore slightly collapsed showing the three rays along one of which the spore has
burst, x 400. ¥.31517. Sandbanks, Dorset.

FIG. 29. A spore showing the three rays very clearly, x 400. ¥.31513. Branksome
Dene, Dorset.

FIG. 30. A somewhat crumpled spore, x 400. ¥. 31513. Branksome Dene, Dorset.

FIG. 31. A somewhat crumpled detached spore almost certainly of this species, x 400.
¥.31518. Alum Bay, Isle of Wight.

Bull. B.M. (N.H.) Geol. 2, 1

PLATE 33

sw

ANEMIA POOLENSIS, A. SCHIMPERIANA (Recent)

PLATE 34

Anemia poolensis n. sp.

FIG. 32. Microtome section through a pinnule segment showing more than one layer of
thin- walled cells, x 150. Section now disorganized. ¥.31509.

FIG. 33. Microtome section across a recurved fertile pinnule. It shows the overlapping of
pinnule segments, sporangia packed with spores close together on the lower surfaces of the seg-
ments, and the thick-walled cells of annuli usually in section, x 50. V. 31510.

FIG. 34. Part of another section across the same pinnule. Two overlapping segments are
shown, many cells thick in this part of the pinnule. One sporangium in section and parts of
two others, one with annulus displayed, x 150.

FIG. 35. Another microtome section showing the annuli and spores of two sporangia, x 150.
Section now disorganized. ¥.31509.

FIG. 36. Microtome section showing thickened tracheids in the pinnule segment. Part of an
annulus is seen below, x 400. ¥.31506.

FIG. 3.7. Another section of the same pinnule showing overlapping pinnule segments, many
cells thick proximally, thinning to a few cells thick only, distally. A partly collapsed paraphysis
(p) is seen arising from one of the segments. Two sporangia are visible in section, that on the
right showing much of the annulus. Below it is a small patch of chloroplast granules (chl)
probably inside one of the cells of the sporangium wall. Spores are visible in the sporangia.
X 150. ¥.31506.

FIG. 38. Part of the same, enlarged, to show the group of chloroplasts just below the thick-
walled cells of the annulus. x 400.

All the above are from Lake, Dorset.

Bull. B.M. (N.H.) Geol. 2, 7

PLATE 34

-^ ~* • "*" ^ tfl

PU*> ,<~~*L

ANEMIA POOLENSIS

GEOL. n, 7. 27

PLATE 35

Anemia colwellensis n. sp.

FIG. 39. Annulus, imperfect, from above, x 150. ¥.31519.

FIG. 40. Part of another annulus, torn and distorted, from above, x 150. V. 31520.

Anemia poolensis n. sp.

FIG. 41. Annulus, from above, for comparison. The cells which have the appearance of an
apical plate are probably a displaced fragment of sporangial wall. Each shows one dark spot
(chloroplast?). x 94. ¥.31514. Branksome Dene, Dorset.

Anemia colwellensis n. sp.

FIG. 42. Upper half of a sporangium with apical annulus, side view. Two or three cells o:
the apical plate remain. The cells of the sporangium wall immediately below the annulus show
numerous opaque grains of pyrites, perhaps pyrites replacements of chloroplasts. x 94. ¥.31521

FIG. 43. An enlargement of the same, (ap) cells of apical plate, x 150.

FIG. 44. A somewhat distorted sporangium, side, showing the apical annulus above, and the
torn sporangium wall below, x 150. ¥.31520.

FIG. 45. Remains of two sporangia (sp i, sp 2) seen from the side. In both the annulus is
dense and black, their cells being almost indistinguishable. The thinner sporangium wall is
preserved below the annulus in one specimen (sp 2) only. A few pyrites grains (chloroplasts?]
are seen. Spores are visible at (s, s) . x 150. ¥.31522.

FIG. 46. An annulus, from above. The ring of cells is ruptured and the apical plate
destroyed. A spore is seen in the gap. x 150. ¥.31520.

FIG. 47. A paraphysis (p) still attached to a fragment of leaf, x 400. ¥.31520.

FIG. 48. Another isolated paraphysis. x 400. ¥.31520.

FIG. 49. A spore showing the ridges, x 200. ¥.31522.

FIG. 50. The same (partly reflected light) showing the straight untubercled character of the
ridges, x 400.

FIG. 51. The same (transmitted light) focused to show the triradiate mark, x 400.

FIG. 52. The same focused to show spore outline and thickness of wall, x 400.

FIG. 53. Another spore, side view, x 400. ¥.31522.

All the above, except Fig. 41, are from Colwell Bay, Isle of Wight.
All specimens of Anemia colwellensis are from the holotype.

Bull. B.M. (N.H.) Geol. 2, 7

PLATE 35

46

ANEMIA COLWELLENSIS, A. POOLENSIS

PLATE 36

Anemia poolensis n. sp.

FIG. 54. A spore showing the smooth outline and triradiate mark with one forked distal end.
Another spore is seen obscurely out of focus behind, x 400. V. 3 1508.

FIG. 55. Another spore showing outline and smooth surface, x 400. On this slide en-
trapped among the hairs of Anemia poolensis is a single spore of Lygodium poolensis. V.3i5ii.

FIG. 56, 57. Two similar spores, x 400. ¥.31508.

All the above are from Lake, Dorset (Extracted from pinnules).

FIG. 58. A spore showing obscurely the triadiate mark, x 400. ¥.31515. Branksome
Dene, Dorset.

Anemia colwellensis n. sp.

FIG. 59. A spore somewhat collapsed showing ridged surface, x 200. ¥.31522.

FIG. 60. The same spore, x 400.

FIG. 61. Another spore showing the " beaded " effect where the ridges on two surfaces cross
one another, x 400. ¥.31522.

FIG. 62. Another spore showing outline and thickness of wall. It has gaped along two rays
of the triradiate mark, x 400. ¥.31523.

FIG. 63. Another showing a ridge branching and the rather obscure triradiate mark, x 400.
¥.31522.

FIG. 64. Another, side elevation, showing ridges obscurely. The spore has partly folded and
collapsed, x 400. ¥.31523.

All the above are from Colwell Bay, Isle of Wight.

Anemia sp.

FIG. 65. Two impressions of barren pinnules. The nervation is comparable with that seen
on the fertile pinnule of A. colwellensis. x 2. ¥.17498. Thorness Bay, Isle of Wight.
Also figured Reid & Chandler, 1926, pi. i, fig. 11.

Lygodium prestwichii Gardner

FIG. 66. Impression of the upper surface of a bilobed pinnule, one lobe well exposed, the other,
rolled up, at lz. The toothed margin is seen on the left of the larger lobe. The secondary nerves,
sunk in the original, appear as fine thread-like ridges, the primary nerve appears as a sharp
ridge in the impression representing a sharply angled channel in the original. x 2. ¥.24862.
Newington, Kent.

FIG. 67. The counterpart of the above showing the impression of the lower surface with
conspicuous rounded midrib in the original (a rounded channel in the fossil) and the raised thread-
like impressions of the sunk secondary nerves of the original. Second lobe of leaf at / 2. x 2.

FIG. 68. The teeth on the right margin in Fig. 67. The thread-like secondary nerves are
also seen and the thickening of the margin, x 6-5.

FIG. 69. An impression of the lower surface of another pinnule with well-preserved nerves
but poorly preserved margins scarcely showing the teeth, x 2. ¥.24859. Newington, Kent.

Bull. B.M. (N.H.) Geol. 2, 7

PLATE 36

m

ANEMIA POOLENSIS, A. COLWELLENSIS, ANEMIA SP., LYGODIUM PRESTWICHI

PLATE 37

Lygodium kaulfussi Heer

FIG. 70. Impression of a barren-lobed pinnule showing nervation and entire margin, x 1-8.
V.2865Q.

FIG. 71. Another long narrow pinnule lobe, x 1-8. V. 28658.

FIG. 72. Fertile pinnule, upper surface, x 15-5. ^.31524.

Fig. 73. The same, lower surface, x 15-5.

FIG. 74. Another fertile pinnule (incomplete at the base), upper surface, x 15-5. ^.31525.

FIG. 75. The same, lower surface. It shows clearly the marked sinuosity of the midrib.

x 15-5-

FIG. 76. A distorted fertile pinnule showing the long stalk, x 15-5. V. 31526.

FIG. 77. The same, opposite side, x 15-5.

Fig. 78. A fertile pinnule, upper surface, somewhat battered and sand pitted, x 15-5.
Specimen destroyed in macerating to release few remaining spores (slide V. 31 531). Branksome
Dene, Dorset.

FIG. 79. Cuticle fragment from a fertile pinnule showing sinuous cells and " beaded " thicken-
ing, giving a " dotted " effect, x 150. ¥.31527.

FIGS. 80-83. Typical spores extracted from two fertile pinnules, showing the smooth surface
and triradiate mark. Folds shown are due to partial collapse, x 400. Figs. 80, 81 are from
¥.31528. Figs. 82, 83 are from ¥.31529.

All the above, except Fig. 78, are from Studland, Dorset.

Bull. B.M. (N.H.) Geol. 2, 7

PLATE 37

LYGODIUM KAULFUSSI

PLATE 38

Lygodium kaulfussi Heer

FIG. 84. An unusually large and thick-walled spore extracted from one of the fertile pinnules.
It has burst along the triradiate mark, x 400. V. 3 1530. From the pinnule which yielded the
spores figured in PI. 37, figs. 82, 83 (¥.31529).

FIG. 85. A typical spore showing the smooth surface and triradiate mark. Folds are due to
collapse, x 400. ¥.31528. The spores in PI. 37, figs. 80, 81 are from the same pinnule.

Both the above are from Studland, Dorset

FIG. 86. A large spore removed from the fertile pinnule (PI. 37, fig. 78) showing the smooth
surface and triradiate mark and small fork at the termination of the ray at the top of the figure.
X 400. ¥.31531. Branksome Dene, Dorset.

FIG. 87. A collapsed isolated spore apparently of this species, x 400. ¥.31532. Cliff
End, nr. Mudeford, Hampshire.

Lygodium poolensis n. sp.

FIG. 88. A naturally macerated fragment of fertile pinnule showing the recurved sporangia
exposed by the decay of the enveloping pinnule segments and bracts. Only one (x) is complete.
This one shows the annulus by reflected light, r, r remains of other sporangia. x 50.
¥.31533. Holotype.

FIG. 89. Diagram of the sporangium (x) in Fig. 88 to show the position of the annulus which
is very obscurely seen in the photograph.

FIGS. 90-96. Four spores from the above fertile pinnule. They show the deeply pitted surface
(the pits bounded by the walls of confluent tubercles?). Figs. 95, 96 represent the same crushed
spore. Fig. 95 is focused to show the thickness of the wall, Fig. 96 the extent of the triradiate
mark. Figs. 90, 91 and 94 also show the triradiate mark. Figs. 92-94 represent the same spore
differently focused to show triradiate mark, outline and surface ornamentation. Pits show best
in Fig. 93, tubercles forming the walls of pits in Fig. 92 (top right). All x 400. ¥.31534-36.

All the above are from Lake, Dorset.

Lygodium scandens Sw. (Recent)

FIGS. 97-101. Spores. Fig. 97 shows a crushed spore which has split along the triradiate
mark. Fig. 98, the triradiate mark in an uncrushed spore. Fig. 99 is the same as Fig. 98 but
the focus shows the wall thickness and pitted surface. Fig. 100, another spore showing surface
structure and pitting. Fig. 101, another focused to show wall thickness in optical section.
It also shows obscurely the triradiate mark. Figs. 97, 99-101. x 400. ¥.31537. Fig. 98.
X 400 approx. Recent (Ross 123. Nigeria).

Bull. B.M. (N.H.) Geol. 2, 7

PLATE 38

96

98

101

LYGODIUM KAULFUSSI. L. POOLENSIS. L. SCANDENS (Recent)

1 OCT 1955

PRINTED IN GREAT BRITAIN BY
ADLARD AND SON, LIMITED
BARTHOLOMEW PRESS, DORKING

r,,

•>)

1956 THE BRACHYOPID

LABYRINTHODONTS

D. M. S. WATSON

BULLETIN OF
THE BRITISH MUSEUM (NATURAL HISTORY)

GEOLOGY Vol. 2 No. 8

LONDON: 1956

BULLETIN OF
THE BRITISH MUSEUM (NATURAL HISTORY)

GEOLOGY

Volume I (1949-52) :

Price
No. i (1949). The Pterobranch Rhabdopleura in the English Eocene.

H. D. Thomas & A. G. Davis . 75. 6d.

No. 2 (1949). A Reconsideration of the Galley Hill Skeleton. K. P.

Oakley & M. F. Ashley Montagu ...... 55.

No. 3 (1950). The Vertebrate Faunas of the Lower Old Red Sandstone

of the Welsh Borders. E. I. White

Pteraspis leathensis White a Dittonian Zone-Fossil. E. I.

White 75. 6d.

No. 4 (1950). A New Tithonian Ammonoid Fauna from Kurdistan,

Northern Iraq. L. F. Spath ....... 105.

No. 5 (1951). Cretaceous and Eocene Peduncles of the Cirripede Euscal-

pellum. T. H. Withers 55.

No. 6 (1951). Some Jurassic and Cretaceous Crabs (Prosoponidae) .

T. H. Withers 55.

No. 7 (1952). A New Trochiliscus (Charophyta) from the Downtonian

of Podolia. W. N. Croft 105.

No. 8 (1952). Cretaceous and Tertiary Foraminifera from the Middle

East. T. F. Grimsdale 105.

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Volume II (1953-56) :

No. i (1953). The Deer of the Weybourn Crag and Forest Bed of

Norfolk. A. Azzaroli ........ 255.

No. 2 (1953). A Coniferous Petrified Forest in Patagonia. M. G.

Calder ........... 125.

No. 3 (1953). The Solution of the Piltdown Problem. J. S. Weiner,

K. P. Oakley & W. E. Le Gros Clark 35. 6d.

No. 4 (1954). Some Upper Cretaceous and Eocene Fruits from Egypt.

M. E. J. Chandler i6s.

No. 5 (1954). The Carboniferous Flora of Peru. W. J. Jongmans . 155.
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Problem. J. S. Weiner, W. E. Le Gros Clark & K. P. Oakley

et al. . . . . . . . . . . 2os.

No. 7 (1955). The Schizaeaceae of the South of England in Early

Tertiary Times. M. E. J. Chandler . . . . . . 155.

No. 8 (1956). The Brachyopid Labyrinthodonts. D. M. S. Watson . 20.?.

THE BRACHYOPID LABYRINTHODONTS

BY

DAVID MEREDITH SEARES WATSON, F.R.S.

Trustee of the British. Museum

Pp- 315-392 ; PL 39 i 34 Text-figures

BULLETIN OF
THE BRITISH MUSEUM (NATURAL HISTORY)

GEOLOGY Vol. 2 No. 8

LONDON : 1956

THE BULLETIN OF THE BRITISH MUSEUM
(NATURAL HISTORY), instituted in 1949, is
issued in five series corresponding to the Departments
of the Museum, and an Historical series.

Parts appear at irregular intervals as they become
ready. Volumes will contain about three or four
hundred pages, and will not necessarily be completed
within one calendar year.

This paper is Vol. 2, No. 8 of the Geological series.

PRINTED BY ORDER OF THE TRUSTEES OF
THE BRITISH MUSEUM

Issued February, 1956 Price Twenty Shillings

THE BRACHYOPID LABYRINTHODONTS

By D. M. S. WATSON

CONTENTS

(1) INTRODUCTION ............

(2) BRACHYOPIDS PROPER ...........

(i) Survey of group ...........

(ii) Bothriceps australis ..........

(iii) B. major — Trucheosaurus gen. nov. ........

(iv) Brachyops laticeps ...........

(v) Batrachosuchus watsoni ..........

(vi) Batrachosuchus browni ..........

(vii) ? Batrachosuchus sp. ..........

(viii) Pelorocephalus mendozensis .........

(ix) " Platyceps " wilkinsoni ..........

(3) " PLAGIOSAURIDS " ...........

(i) Description of material ..........

(ii) Points of agreement with older Brachyopids ......

(iii) Dvinosaurus compared with older Brachyopids .....

(4) EOBRACHYOPS ............

(i) Description of material and manner of fossilization .....

(ii) Skull roof

(iii) Occiput ............

(iv) Palate

(v) Lateral line ............

(vi) Lower jaw ............

(vii) Vertebral column ...........

(viii) Shoulder girdle ...........

(ix) Scales .............

(x) Possible relationship to Acheloma casei and Trimerorhachis ....

(5) TRIMERORHACHIS ............

(i) Variation of specimens ..........

(ii) Brain-case ............

(6) COMPARISON OF EOBRACHYOPS WITH TRIMERORHACHIS .....

(i) Resemblances ...........

(ii) Differences ............

(iii) Two striking common features ........

(iv) Characteristics of Eobrachyops are the special qualities of Brachyopids .

(v) Conclusion on relationship of Eobrachyops to Trimerorhachis and Brachyopids

(7) EOBRACHYOPS COMPARED WITH PELION ........

(i) Identification of Pelion lyelli .........

(ii) Re-examination of Pelion .........

(iii) Probable relationships of Pelion ........

GEOL. II, 8,

3i8 THE BRACHYOPID LAB YRINTHODONTS

Page

(8) EOBRACHYOPS COMPARED WITH ERPETOSAURUS ...... 370

(i) Description of Erpetosaurus material ....... 370

(ii) Brain-case ............ 371

(iii) Palate ............. 374

(iv) Vertebral elements . . . . . ... . . -375

(v) Similarities between the two forms . . . . . . -377

(9) PLATYSTEGOS LORICATUM .......... 377

(i) Description of material . . . . . . . . . • 377

(ii) Structure of skull ........... 377

(iii) Comparison with Dendverpeton and Eugyrinus ...... 382

(iv) Comparison with Brachyopids ......... 383

(10) SURVEY OF BRACHYOPID LINE ......... 384

(11) REFERENCES ........... ... 389

INTRODUCTION

THE amphibia now known as " Labyrinthodonts " were first mentioned in print in
G. F. Jaeger (1824 : 10), who, in 1828, published figures of three teeth called Mastodon-
saurus, two " vertebrae " (which are intercentra), and a pair of exoccipitals held
together by parasphenoid which he named " Salamandr aides giganteus ". Later he
recognized that teeth and skull fragments were from the same animal, whose correct
name was Mastodonsaurus giganteus.

In 1841 R. Owen obtained a tooth of Mastodonsaurus and showed (1842) that in
section it had a dentine of unique complexity, found also in fragments of teeth from
Warwick. He called both materials " Labyrinthodon ". This generic name is an
absolute synonym of Mastodonsaurus, but it may well be retained in the name of a
group of higher rank.

During this decade Labyrinthodonts were described in considerable numbers from
the Permian and Trias of Germany and Russia, and the next ten years added Canada,
India, Australia and South Africa to their known distribution, and extended it
downward to the Coal Measures. Thus by 1918 not only had a very large number of
Labyrinthodonts been named, but other groups of Palaeozoic amphibia had been
established, and a great number of new animals attributed to one or other of them.
But the one hundred and twenty-three genera listed in the third edition of Zittel's
Grundzuge by Broili were scattered amongst a number of major groups without real
evidence, and nothing was known of the evolutionary history of any of them. The
best summary was that by O. Abel in Die Stdmme der Wirbeltiere, 1919.

In 1912 I gave a short account of some features of the skulls of two Labyrinthodonts
from the Middle Coal Measures showing that the two, though clearly not closely
related, agreed in many fundamental characters, including a single basioccipital
condyle, reptile-like basipterygoid processes, pterygoids which so closely approach
the middle line as to leave only slits as interpterygoid vacuities, and a narrow
parasphenoid sheathing closely an interorbital septum. These qualities are largely
those of Seymouria, and of Reptiles, but they are also found in the Osteolepid fish
Megalichthys,

THE BRACHYOPID LABYRINTHODONTS 319

In 1919 I reviewed the Permian and Triassic Labyrinthodonts of the world and
showed that they had vertebrae which had separate neural arches, pleurocentra and
intercentra in Rhachitomi — the pleurocentra vanishing in certain late forms, the
" Stereospondyli ". However, the pattern of the dorsal surface of the skull, the
lower jaw, clavicular arrangement, scapulo-coracoid, pelvis and proximal limb bones
do not enable an animal to be referred to one or other group, though they are diag-
nostically Labyrinthodont.

Thus I was enabled to take the great mass of Labyrinthodonts of Permian and
Triassic time together, and by comparing all known Lower Permian with all known
Upper Triassic forms it became clear that the Labyrinthodonts display the following
"advances" :

(1) A gradual flattening of skull and body.

(2) A gradual reduction of the basioccipital and its retreat from the condyle.

(3) A reduction of the basipterygoid articulation and its functional replacement by
a sutural attachment of an enlarged parasphenoid to the pterygoid.

(4) The exoccipitals extend to form the whole occipital condyle, reach forward to
the pterygoids below the ear, and up to sutures with the new occipital flanges of the
dermosupraoccipitals and tabulars.

(5) Reduction and final disappearance of cartilage bones in the paroccipitals,
prootics, supraoccipital, as well as in the basisphenoid and basioccipital.

(6) An exaggeration of the interpterygoid vacuities and a reduction and shortening
of the palatal ramus of the pterygoid.

(7) A reduction of the suspensory part of the skull so that the quadrate condyles
come to lie in front of the occipital condyle.

Subsequently, in 1926, I was able to describe good material of several large
Coal Measure Labyrinthodonts belonging to the two groups of Loxommids and
Anthracosaurs. These animals, although they evidently differ a good deal from
one another, agree in many features ; they have a single nearly circular occipital
condyle to which the small exoccipitals contribute ; the supraoccipital is bony ;
there are no occipital flanges of the tabulars and dermosupraoccipitals ; the basi-
sphenoid has small basipterygoid processes, with articular faces for the pterygoid
(and epipterygoid) ; the parasphenoid has a narrow ramus cultriformis sheathing the
lower edge of an interorbital septum ; the pterygoids meet one another in front, and
leave only narrow slits as interpterygoid vacuities ; the quadrate condyles lie far
behind the occiput. In fact in all these respects they represent a logical starting
point for the series of evolutionary changes deduced from observation of the then
known later Labyrinthodonts. And they evidently make a real approach to the
structure found in Osteolepid fishes.

At that time it seemed obvious that further progress required the adequate descrip-
tion of many other Labyrinthodonts, often named but not giving evidence about
matters of importance. Dr. M. C. Steen therefore began a study of the faunas of fossil
amphibia from the Coal Measures of Nyirany, Bohemia ; Linton, Ohio ; and the
erect trees of South Joggins, Nova Scotia. In a series of papers 1931-1938 she
revised these faunas, immensely increasing our real knowledge of them. Simultan-

320 THE BRACHYOPID LAB YRINTHODONTS

eously A. S. Romer independently revised the Linton fauna, his results agreeing
generally with those of Dr. Steen.

These re-examinations recognized in the Linton fauna a large number of animals
with an incomprehensible vertebral column, which were referred to the Phyllo-
spondyli, together with an obvious Loxommid, and Leptophractus, not yet adequately
known. When in 1937 Dr. Steen examined the Lower Permian Acanthostoma, which
is certainly a Labyrinthodont, she discovered that it possessed paired pleurocentra
and intercentra, which were directly comparable to those of the Linton " Phyllo-
spondyli ", and referred them to the Labyrinthodontia as " Rhachitomi ". Thus the
rhachitomous stage of Labyrinthodont evolution was shown to have occurred in Coal
Measure rocks in Europe and North America, but at a horizon clearly considerably
higher than that of the Newsham and Scottish Coal Measure localities which produced
Loxommids and Anthracosaurs. Finally Dr. Steen described a series of small
closely related Labyrinthodonts from South Joggins of an age which is considerably
older than that of the Linton and NCrany faunas. It is, however, to be noted that
Eugyrinus wildi (A. S. W.), which I held to be a Branchiosaur, is now regarded by
Romer as a " Rhachitomous " Labyrinthodont. Its skull is in fact, though only
18 mm. in length, very much like that of the Dendrerpetons (cf. Text-figs. 29-31 of
Platystegos). Indeed it is the only other form to show the peculiar arrangement
whereby the squamosal is attached to the supratemporal by a special inwardly
directed process. It has enlarged interpterygoid vacuities, relatively larger than those
of Dendrerpeton, and is the oldest Labyrinthodont (or indeed amphibian) to show
them. It has much cartilage bone and is presumably adult.

In 1931 G. Save-Soderbergh, at the age of 21, collected from rocks in East Green-
land, then held to be of Upper Devonian age and certainly containing fish of
Old Red Sandstone type, a series of skulls of amphibia of entirely unknown type.
He published a preliminary description of these fossils in 1932 and called them
the Ichthyostegids. Further materials were collected in later years, but the
prolonged illness and death of Save-Soderbergh prevented any further preparation
and description of them. At last in 1952, twenty years after the original description,
Dr. Jarvik gave us an admirable account of the tail and hind limb of Ichthyostega, a
set of new restorations of the skull of that animal, and a short account of a new,
related, though very different animal — Acanthostega.

Thus it was clear that the whole group of Labyrinthodonts deserved re-examination.
The general scheme of evolution I had described was based on gross comparisons of
faunas of successive age, and it was obviously necessary to attempt to trace individual
evolutionary lines through as long periods as possible, and in considerable detail.
This I had begun, but was unable to complete, so that the appearance in 1947 of a
great review of the Labyrinthodontia by Professor A. S. Romer was most valuable.
Romer made no formal classification, nor did he attempt to isolate individual
evolutionary lines, but he brought the small Labyrinthodonts of Linton, Nyfany,
and elsewhere into his account in an admirable and illuminating manner.

The present paper is an attempt to isolate a single evolutionary group, long lived,
elaborate and world wide, but always recognizable by special structural details found

THE BRACHYOPID L AB Y RINTHODONTS 321

nowhere else. I have work in progress, in a relatively advanced stage, on the
immense material of the Capitosaur series.

In 1915 Robert Broom examined the British Museum type skulls of Owen's
Brachyops laticeps (1854) an<^ Huxley's Bothriceps australis (1859). He had himself
described Batrachosuchus browni (1903), and now pointed out the resemblance
between these three animals, founding for them a new family — Brachyopidae —
without giving any distinctive characters of the group.

In 1919 I described the structure of the posterior part of the palate of Bothriceps,
and gave a more complete account of the skull of Batrachosuchus. With these forms
I brigaded Jaekel's genus Plagiosaurus (1914), founded for P. depressus from Halber-
stadt, to which he had added Fraas's Plagiosternum pulcherrimum (1913) and Fraas's
Plagiosternum granulosum (1889), Jaekel having made for these three forms an
" order " Plagiosauri, without giving any definition of the group.

I defined the group by saying that its members resemble one another, and differ
from all known Stereospondyl types, in the following ways :

(1) The broad parabolic skulls with large anteriorly situated orbits.

(2) The unusual way in which the prootic flange of the squamosal wraps round the
outer side of the quadrate, and having formed a laterally concave face on the occipital
surface, ends in a ridge, separated from an exactly similar ridge of the pterygoid
by the quadrate.

(3) The upturning of the lateral wings of the pterygoids from the subtemporal fossae,
so that the palate forms a broad fl -shaped arch.

(4) The unusual way in which the posterior edge of the pterygoid is applied to the
inner face of the quadrate.

(5) The occipital condyles lie far behind the dermosupraoccipitals so that the
occipital surface slopes forward.

I pointed out that the then undescribed Dvinosaurus possessed all these five
qualities, but was a primitive rhachitomous form. No subsequent author (except
Sushkin, 1936) appears to have paid any attention to the list of diagnostic characters
of the Brachyopids set out above, perhaps because I did not publish a series of
drawings illustrating the matter.

Subsequently Dvinosaurus was described and figured by Amalitzky (1924), by
Sushkin (1923, 1936), and by Bystrow (1935, 1938), the last two bringing out the
neoteny which it shows, and the last producing an admirable account of the whole
skeleton.

The later Brachyopids have been described and discussed by von Huene (1922)
and Tage Nilsson (1934, '37, '39, '45), and A. S. Romer has treated them in his
Review of the Labyrinthodontia (1947). He accepts the Brachyopidae and the group
Plagiosauridae for the extreme forms, but adds the Metoposaurs to the assemblage,
whilst removing Dvinosaurus to the neighbourhood of Trimerorhachis and " Saurer-
peton ". There is thus some variety of opinion about the whole matter and further
treatment of it is necessary.

The Brachyopids proper are rare animals known usually from the occurrence of
single individuals of each species from rocks of Upper Permian and Lower Triassic
age in Australia, India, South Africa and Argentina, and the published accounts of
their structure are inadequate.

322 THE BRACHYOPID LAB YRINTHODONTS

BRACHYOPIDS PROPER
SURVEY OF GROUP

The Brachyopids include a terminal group of Middle and Upper Triassic forms,
the Plagiosauridae, but its typical members are from earlier beds whose age relations
are unfortunately not very accurately known. The complete list of these is as
follows :

(i) Br achy ops laticeps Owen. The type — and only known specimen — was found
in the " Mangali beds " at Mangali, in Nagpur, Central India. This horizon is assumed
on poor evidence to be equivalent to the Panchets (C. S. Fox, 1931 : 159). The age
of the Panchets is to be determined from their fauna of vertebrates, and from a
considerable flora. The fauna has recently been described by von Huene (1942) who
recognizes in it a genuine Lystrosaurus and remains of Chasmatosaurus , genera
otherwise known only in the Lystrosaurus zone of South Africa and of Sinkiang.
Furthermore he has described a skull of the Labyrinthodont Gonioglyptus , which
genus occurs in the Panchets, from the Prionolobus beds of Chideru in the Salt
Range. I am not, however, convinced that the generic identity of the two animals
is established. The important member of the flora is Glossopteris , which seems to be
the commonest form, and is accompanied by its stem " Vertebra ria ", so that the
generic determination may be regarded as certain. It is primarily a Permian form,
found in South Africa from the Ecca upward, though it probably does not occur in
the Cynognathus zone flora of Aliwal North. In Australia it is abundant from the
Lower Marine series up through the Coal Measures of Newcastle, but it probably
does not occur in the Narrabeen shales — or higher. Thus it can probably be assumed
that Br achy ops is of Lystrosaurus zone age.

(ii) Bothriceps australis Huxley. The type — and only known specimen — was
bought by the British Museum in 1848 from a person of whom nothing is known, and
was then said to have been found in " Australia ".

(iii) "Platyceps" wilkinsoni Stephens. There were several described specimens
from the well known fish bed of Gosford, New South Wales, which lies in a sandstone
member of the Narrabeen shales. One specimen was on loan to the British Museum
(Natural History) about 1920-24, when I had a photograph made from it. Its age
is relatively well determined ; it is in a formation immediately followed conformably
by the Hawkesbury sandstone, from which came the well-preserved fish fauna of
Brookvale, described by R. T. Wade in 1935. This fauna is directly comparable with
that from Bekker's Kraal in the Cynognathus zone of South Africa, and thus appears
to be Lower Triassic.

(iv) "Bothriceps " major A. S. W. (non Owen) comes from a " coal " — actually a
torbanite — at Airly, Central Coalfield, New South Wales. This lies in the " Upper
Coal Measures ", a formation succeeded by the Narrabeen shales. It is therefore of
pre-Triassic age, and may be nearly contemporary with Brachyops, or perhaps a
little older, in the definitely Permian Cistecephalus zone.

(v) Batrachosuchus browni Broom comes from Aliwal North, Cape Province,
South Africa, presumably from the Cynognathus zone, Lower Trias.

THE BRACHYOPID LAB YRINTHODONTS 323

(vi) Batrachosuchus watsoni Haughton comes from an unknown locality certainly
in the Cynognathus zone of the Burghersdorp district.

(vii) Pelorocephalus mendozensis Cabrera comes from rocks at Potrerillos, Mendoza,
Argentina, which on inadequate evidence are regarded as of Upper Middle Triassic
age.

Thus in order of age the Brachyopids are :
Rhaetic — Gerrothorax, Plagiosaurus.
Upper Trias — Gerrothorax, Plagiosternum, Plagiosuchus .
Middle Trias — Plagiosuchus, ? Pelorocephalus.
Lower Trias — Batrachosuchus, Platyceps.
Basal Trias, or Uppermost Permian — Br achy ops.
Upper Permian (Cistecephalus zone) — "Bothriceps " major.

Its own structure suggests that Bothriceps australis is as early as, or earlier than
"Bothriceps " major. Dvinosaurus from the Cistecephalus zone of Russia is presum-
ably of much the same age as "Bothriceps " major, and may well have been contem-
porary with Bothriceps australis, or indeed younger.

BOTHRICEPS AUSTRALIS Huxley

The unique type specimen (B.M.N.H., 23110) consists of a skull with the lower jaw
tightly closed on it. The whole upper and lateral surface is preserved as a beautiful
internal cast, showing the details perfectly, except for some lost regions. To this
cast a little actual bone adheres in places and shows, perfectly preserved, the outer
surface over small areas, including a lachrymal. The occiput, though damaged,
remains as well-preserved bone, and the palate (further developed since 1919) is
excellently preserved. Both lower jaws are complete and well shown.

The skull is parabolic in plan and about 15% wider than its length, measured on
the skull table. On the left side of the specimen it can very clearly be seen that as
far back as the orbit the anterior part of the skull is very much depressed, its ventral
border then turning downward so that the skull becomes very deep at the occipital
border. The orbits lie almost entirely in front of the mid length, and are compara-
tively small, so that the face is long for a Brachyopid. The hinder border of the
dermal roof is damaged at the lateral end of the tabular, but it seems obvious that
there can have been no real otic notch ; indeed there may not have been any
embayment at all. The dermosupraoccipitals have a well-marked posterior border
on the upper surface, behind which they are produced downward on to the sloping
occipital surface by lappets which rest on and end in a suture with the exoccipitals.

The tabulars are short but wide, having a long suture with the supratemporal
and the squamosal. They have long occipital flanges passing downward and in-
ward below the large post-temporal fossae to meet the exoccipitals in suture.

The shapes and relations of the main bones of the skull roof can best be understood
from Text-figs. I and 2. The more important features to which attention should be
drawn are : there is a septomaxilla, clearly shown as a small triangular area of bone
surrounded by sutures, which is attached to the nasal and lachrymal behind the
nostril. The lachrymal is shown on both sides of the skull ; in part the sutures

324

THE BRACHYOPID LABYRINTHODONTS

For. Maa.

Tab. D.S.Oc. p.T.Fos.

TEXT-FIG, i. — Bothriceps australis Huxley. Type specimen (B.M.N.H., 23110) slightly
restored. Natural size. A, Dorsal surface ; B, occipital view, with lower jaw in place.
The detailed shape of the bones on the skull roof comes nearly entirely from the impres-
sion of their under surface, thus the fragments of lateral line grooves shown on the outer
surface of the right premaxilla, left lachrymal, and right jugal and squamosal are not
represented. Ang., angular ; Art., articular ; D.S.Oc., dermosupraoccipital ; Ex.Oc.,
exoccipital ; For. Mag., foramen magnum ; Fr., frontal ; Ju., jugal ; Lac., lachrymal ;
MX., maxilla ; Na., nasal ; P. MX., premaxilla ; P.O., postorbital ; P.T.Fos., post-
temporal fossa ; Par., parietal ; Pr.Art., prearticular ; Pr.Fr., prefrontal ; Pt.,
pterygoid ; Pt.Fr., postfrontal ; Qu., quadrate ; Qu.J '., quadratojugal ; S.Mx., septo-
maxilla ; S.Oc., supraoccipital ; S.Tem., supratemporal ; Sq., squamosal ; Sur.Ang.,
surangular ; Tab., tabular ; X, tenth cranial nerve foramen.

3*5

Vo

Pol.

Art

Ex.Oc

Ex

Lac.

Art.

P.Mx.

S.Ang.

TEXT-FIG. 2. — Bothriceps australis Huxley. Natural size. A, Ventral surface, showing
imaginary removal of the hinder part of the right mandible ; B, right lateral aspect,
showing ornament and lateral line groove where preserved. The lower jaw is restored
to its natural position by being lowered so that the quadrate articulated with it regains
its original place, and the ornamented outer surface of the angular does not underlie
the quadratojugal. The projection of the dentary in front of the premaxilla and the
backward direction of its anterior teeth are as in the specimen. Reference letters as
before with : Den., dentary ; Pal., palatine ; Par.Sp., parasphenoid ; Vo., vomer ;
VII, foramen for seventh cranial nerve (chorda tympani).

326 THE BRACHYOPID LABYRINTHODONTS

surrounding it are shown on the outer surface of the bone, elsewhere on the mould of
the inner surface. The bone does not enter the margin of the nostril, but it is
possible that it does form a very small part of the orbital margin. It is entirely
surrounded by the maxilla, septomaxilla, nasal, prefrontal and jugal. The pre- and
postf rentals meet in suture above the orbit. The jugal forms the whole lateral
margin of the orbit, articulating in front with the pre-frontal ? and lachrymal.
The large supratemporal meets the postfrontal and postorbital anteriorly, and the
dermosupraoccipital and tabular behind. The squamosal is represented mainly by an
internal impression, but some bone is preserved. It has a large superficial exposure
of ornamented bone ending behind at a definite margin round which the bone turns
on to the occipital surface. Here it forms a smooth face standing nearly vertically,
and concave on a vertical axis, so that its admesial border is turned backward ending
parallel with, but separated by a narrow space from, the hinder border of the quadrate
ramus of the pterygoid. This space was obviously occupied by a cartilaginous ridge
arising from the visible occipital surface of the quadrate. The quadratojugal is a
relatively large bone, lying below and continuing the surface of the squamosal, and
extending forward to meet the jugal. That part of it which lies on the outer surface
has a normal labyrinthodont ornament, the occipital area being smooth.

The occipital surface is preserved in bone, but is much damaged by weathering and
fracture. None the less its structure can easily be made out, and it is represented
in Text-fig. I. The occipital surface slopes backward for a long distance (its own
height) behind the ridge on the dermosupraoccipital which marks the hinder border
of the skull table. There is no bony basioccipital, but a space which must have been
occupied by it exists dorsal to the hinder end of the parasphenoid and between the
roots of the exoccipital condyles. The existence of a cartilaginous supraoccipital is
shown by the usual shelves on the exoccipitals. The exoccipital condyles are broken
off, and the broken surface polished, but they were evidently transversely widened
and had a considerable projection backward. The main body of the bone extends
upward to meet the occipital lappet of the dermosupraoccipital in a suture, then
outward to a suture with the occipital flange of the tabular, forming the lower border
of a post-temporal fossa, and evidently sheathing the posterior surface of a cartila-
ginous paroccipital. The lower part of the bone there passes forward in contact with
the upper surface of the lateral border of the parasphenoid to end in a contact with
the pterygoid. The exoccipital is perforated by a foramen for the vagus, but it is
impossible to be sure whether or not a hypoglossal foramen existed.

The palate, after further preparation by Mr. L. E. Parsons, is well exposed on
the left side of the skull, although the adherent lower jaw hides its lateral portions.
The parasphenoid has a processus cultriformis with a flat ventral surface which
expands into a widened body whose lower surface is also flat, though it is crossed by
two shallow grooves which meet in the middle line. It ends in a shallow notch
between the pedicels of the exoccipital condyles, the bone is then in contact with the
exoccipital by a nearly straight suture extending outward and forward to meet the
long one between the parasphenoid and pterygoid. This suture has a rather charac-
teristic sinuous course which can be seen in Text-fig. 2, A. There is no transverse
ridge corresponding to that which in Capitosaurus represents the pockets for the

THE BRACHYOPID LAB YRINTHODONTS 327

recti capitis muscles. The pterygoid at its attachment to the parasphenoid is flat,
and this horizontal surface extends outward until it rather suddenly turns downward
to end at the margin of the subtemporal fossa. The vertical wall so formed extends
backward to lie in contact with the admesial surface of the quadrate, so that a thin
strip of that bone, and its cartilaginous dorsal continuation, separate it from the
corresponding inner flange of the squamosal. It is evident that the lower part of the
wall was a sheath covering the inner side of the masticatory muscles. The inter-
pterygoid vacuities are large, their anterior margins lying in the palatines and vomers.
There is an alternative pair of large teeth on the palatine, and one very laterally
placed on the vomer. The border of the internal nostril is not shown.

The lower jaw is in postion with the mouth shut. In the specimen the teeth at
the front end of the lower jaw lie in front of those in the premaxillae, and much of
the maxilla lies mesial of the dentary tooth row. How far this condition is natural I
do not know. The lower jaw is shallow and wide. Its ventral surface is straight
and there is a large retro-articular process made from the articular, covered laterally
by the surangular and angular, and mesially by a prearticular which, just anterior
to the articulation with the quadrate, is perforated by a foramen for the chorda
tympani. A small Meckelian foramen between the prearticular and angular lies on
the inner surface just above the ventral border of the jaw.

" BOTHRICEPS MAJOR " A. S. W. = TRUCHEOSAURUS gen. nov.

The type specimen of Bothriceps major, found at Airly in an " oil shale " in the
Newcastle Coal Measures of the Central Coalfield of New South Wales, is a large part
of a skeleton, one surface being on B.M.N.H., R.3728, whilst the counterpart is in
Sydney, New South Wales. The slab in which it is preserved is a bog-head or
torbanite, no doubt made largely of the alga Reinschia. This material was certainly
deposited on the floor of a lake or pool, and the animal must have sunk on to it
whilst still nearly (or quite) complete. It has preserved the bones extremely badly,
little of them remaining except as a buff coloured film of very small thickness. Thus
no preparation is possible.

The specimen in London now consists of two blocks. One contains the head,
broken so that the left hinder corner is lost and the occiput is incomplete. The
other shows a series of 28 vertebrae and their ribs, no trace of the shoulder girdle
remaining. This block extends beyond the preserved part of the vertebral column.

The head and lower jaw are dorsoventrally compressed, and only a faint impression
of the upper surface remaining on the slab can be interpreted. The general shape is
evident, but it is impossible to be sure of the position of the orbits. The almost
straight hinder edge of the dermal roof of the skull is certain, and the lappets of the
tabular and dermosupraoccipital which extend backward on to the sloping occiput
are well shown. The sutures shown in Text-fig. 3 are derived mainty from the abrupt
changes of direction of the ridges which form the ornament of the bones of the skull
roof, and these are in all probability a close approximation to the facts. There are no
visible traces of lateral line grooves. This skull is evidently that of a Brachyopid :
the shortness, great width, presumed anterior position of the orbits, and especially

328 THE BRACHYOPID L A B Y RI NTHODONT S

the absence of an}' otic notch, and straight occipital margin of the skull roof,
with the long projection of the occipital surface behind the roof — all facts which
are clearly shown and certain — are to be found only in this group of Labyrin-
thodonts. A small piece of matrix surrounding the occipital process of the tabular
and the hinder border of the squamosal on the right side remains. Had the specimen
possessed ossified branchial arches some parts of them would have been expected to
be preserved in this area, but as there are no such traces visible it is probable that the
species was not neotenous. Thus the Brachyopids existed in a typical form in the
Upper Permian, presumably in the Cistecephalus zone.

Tab.

D.S.Oc.

TEXT-FIG. 3. — Trucheosaurus (" Bothriceps ") major (A. S. W.) (B.M.N.H.
Diagram of skull roof, x J. Reference letters as before.

The remainder of the skeleton shows little. The vertebrae are so crushed dorso-
ventrally that nothing can be said about them, except that they do not seem to have
had the solidity and close approximation of those of the latter Plagiosuchids. The
ribs are parallel-sided, little curved, and short — consistent with a broad, dorso-
ventrally flattened body. There is evidence of some 28 vertebrae, and their pairs
of ribs, preserved ; the slab containing the shoulder region is likely to have held
5 more ; the series ends before there is any sign of a pelvis and hind legs, which must
have lain even further back, although the preserved and inferred part of the backbone
is three and a half times as long as the skull. A misplaced fore leg shows a humerus
less than a quarter of the skull length in its bony extent, and shorter radius and ulna.

That Bothriceps major A. S. W. is a Brachyopid is thus certain, but in the few
structures clearly shown in the only known specimen it differs from the type of the
genus Bothriceps, and indeed from all other known genera of the family. It is thus
desirable to place it in a new genus, but the specimen is so poorly preserved that it
is quite probable that a well preserved skull of the same species might not be recog-
nizable as such, so that any new genus founded for the specimen would naturally
remain without adequate definition. But the specimen is important as showing the

THE BRACHYOPID LAB YRINTHODONTS

RHAETIC

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Gerrothorax (Sweden fa N-Germany)

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330

THE BRACHYOPID LAB YRINTHODONTS

occurrence of a typical Brachyopid at an early horizon, and it must have a generic
name for reference, for which I propose Trucheosaums (Gr. rpvxea, rags and tatters).

Ex.Oc.Con.

B

Ex.Oc.Con.

TEXT-FIG. 4. — -Brachyops laticeps Owen. Type specimen (B.M.N.H., 11.4414). x J.
A, Direct dorsal view ; B, lateral view, without reconstruction, dotted outline is that
of the matrix. Reference letters as before, with : Ex.Oc.Con., exoccipital condyle. •

BRACHYOPS LATICEPS Owen

The unique type specimen of Brachyops laticeps Owen (B.M.N.H., R.44I4) was
found at Mangali, Nagpur, Central India, in beds of uncertain correlation but of
probable basal Triassic age.

This skull, which lacks a lower jaw, is contained in a curious pink-coloured fine-
grained rock, and has very nearly retained its original shape, though the right side
is slightly crushed downward and outward. The preservation is unusual and bad.
The actual bone seems to have lost its structure ; even when seen in transverse

THE BRACHYOPID LAB YRINTHODONTS 331

section the hinder ramus of the pterygoid is thin and not recognizable under a lens as
bone. The superficial ornament of the upper surface is recognizable, though pre-
served in unnaturally low relief, and imperfectly. The right orbit is well defined,
but the position of the pineal foramen is uncertain. The preorbital part of the skull
shows its outline, and a possible stretch of the border of the nostril, but is otherwise
unintelligible. Sutures are occasionally visible, but must often be inferred from the
radiate ornament of the individual bones. Lateral line grooves are sometimes
evident. The occiput is present, badly preserved The structure of the dermal skull
roof is obvious from Text-fig. 4, A and B, in which the projection of the occiput is
certainly fixed, as is the width across the occipital condyles. The characteristic
straight occipital border and absence of otic notches are certain, and the specimen
shows clearly on both sides the swinging round of the squamosal on to the hinder
surface and its termination at a backwardly turned ridge, clearly seen on the left
side to be separated by a narrow space of nearly constant width from the hinder edge
of the pterygoid seen in section. This space obviously received a cartilaginous ridge
on the posterior surface of the quadrate. The lateral line grooves, whose distribution
is shown, are relatively narrow, unlike those of Batrachosuchus.

In side view (Text-fig. 4, B) it is evident that the hinder part of the maxilla lower
(tooth bearing) border is turned a little downward, the lower border of the jugal and
quadratojugal continuing its trend. (Dinkel's admirable drawings (Owen, 1855) do
not bring this point out clearly, presumably because of some foreshortening by use
of a camera lucida.)

BATRACHOSUCHUS WATSONI Haughton

The genus Batrachosuchus was founded by Broom for B. browni, an excellent skull
found by Alfred Brown in the Cynognathus beds of the Aliwal North district, Cape
Province, South Africa. The only other material is a skull, B.M.N.H., 1^.3589,
from the Seeley collection, which is without any record of origin, but almost certainly
came from the Cynognathus zone of the Burghersdorp district. I gave a description
of this skull in 1919, and S. H. Haughton subsequently (1925) made it the type of
B. watsoni. This skull is nearly complete and most perfectly preserved. It is quite
undistorted and shows all details of the bones in perfection, but unfortunately the
lateral borders of the orbits have been broken away and lost, and the lower parts of
the jugals and quadratojugals have been removed, leaving however an impression of
the inner surfaces which shows the lower border of the skull with certainty. In my
original description I illustrated the skull roof only by a photograph, which was
taken at the British Museum in my absence. This was evidently made with a
vertical camera, the skull resting on the tip of the quadrate, and on a piece of matrix
which supports the premaxillary teeth, when the skull roof lies at an angle of more
than 30 degrees with the horizontal. Thus my published photograph is fore-
shortened, and exaggerates the projection of the occiput and shortness of the skull in
relation to its width.

The pattern of the dermal bones of the skull roof is shown in Text-fig. 5 in which it
is seen at right angles to its plane, and to the sagittal plane of the skull from a distance

GEOL. II, 8. 2

332

THE BRACHYOPID LAB YRINTHODONTS

P. MX.

S.Mx.

MX.

Pr. Fr.

Tab.

Sub.Tem.Fos.

TEXT-FIG. 5. — Batrachosuchus watsoni Haughton. Type specimen (B.M.N.H., 11.3589).
X -f. A, Direct view at right angles to dorsal surface, essentially unrestored ; B,
direct lateral view. The broken line lower border of the temporal region from the
hinder end of the maxilla backward follows an impression of the visceral surface of the
bones, and is certain. (The quadrate, though shown in solid line, is considerably
eroded). Reference letters as before with : Ec.Pt., ectopterygoid ; St., stapes ;
Sub. Tern. Fos., subtemporal fossa,

THE BRACHYOPID LABYRINTHODONTS

333

Pal.

Ec.Pt.

Pt

TEXT-FIG. 6. — Batrachosuchus watsoni Haughton. Type specimen (B.M.N.H., 11.3589).
X -|. The palate, viewed at right angles to the parasphenoid. Reference letters as
before. Qu.J ' . is a transverse section of the quadratojugal where it turns inward behind
the quadrate to form the characteristic concave surface found only in Brachyopids.
It is drawn as it exists in the specimen.

TEXT-FIG. 7. — Batrachosuchus watsoni Haughton. Type specimen (B.M.N.H., 11.3589).
X f . The occiput seen directly from behind with the upper surface horizontal.
Reference letters as before with : B.Oc.Proc., process of exoccipital over the basioccipital
cartilage and below the brain cavity ; Par.Ot.Pt., the parotic part of the pterygoid
as now exposed, it may reach the skull roof, and extend far in.

334 THE BRACHYOPID L AB YRINTHODONTS

of about 100 cm. The only region where the skull roof pattern presents any difficulty
is that between the nostril and the orbit. The structure of the premaxillae is obvious;
they meet in the middle line, a small opening — perhaps for an interpremaxillary
gland — interrupting the suture. They are attached to the anterior ends of the nasals,
the two bones together forming the anterior, mesial and hinder borders of the some-
what asymmetrical nostrils. On each side the margin of the nostril is completed by a
small bone attached by obvious sutures to the lateral part of the premaxilla in front,
and to the lateral part of the nasal behind. These bones are ornamented, each bearing
a small knob on its upper surface anteriorly, that on the left having further ornament
posteriorly. On the left side I excavated the nostril to its floor, exposing the smooth
upper surface of the vomer (separated by a distinct suture from the premaxilla) and
found that the bone now under discussion descends from the outer border of the
nostril to expand into a small flat sheet resting on the upper surface of the vomer.
The posterior end of the vertical part of the bone is notched, and it is clearly a
septomaxilla, a simple modification of the ordinary labyrinthodont type.

The anterior part of the maxilla is perfectly preserved on each side, its lower
border, and the teeth which it bears, being visible on the left side. From here its
very highly sculptured surface can be traced upward to an obvious suture with the
premaxilla, septomaxilla, nasal and prefrontal. There is thus no sign of a lachrymal ;
either the bone has fused early in life with the maxilla — which is improbable — or it
has vanished, squeezed as it were out of existence by the approximation of the
nostril and orbit.

Nothing further in the structure of the skull roof calls for notice here ; the whole
pattern is determined with certainty. The noticeably anterior position of the
quadrates, obscured in my early photograph, is important.

The lateral aspect of the skull (Text-fig. 5, B) brings out its extraordinary shape.
The preorbital region is low, there being in fact only a space of some five millimetres
in height separating the upper surface of the vomer from the lower surface of the
nasal at the hinder margin of the nostril. The depth of the skull at the orbit is much
less than the length of that opening, and unless the eye were very small it must have
projected far above the skull roof, like that of a frog, and, like the frog's, have been
capable of retraction through a special lateral bay in the interpterygoid vacuity. It
is remarkable that the orbital margin is not at all everted as is that of Capitosaurus.
Behind the orbit the skull rapidly deepens until at the quadrate it is remarkably
high. The dorsal surface is flat, but it seems evident that there was a break in the
direction of the ventral border of the skull roof about at the back of the orbit, the
lower margin of the maxilla lying nearly parallel to the skull roof, whilst that of the
jugal and quadrate jugal lies at an angle of some 40 degrees to it. Thus, as shown
in my original figure of the occiput, the quadrate condyle is placed very ventrally,
lying far below the ventral surface of the basis cranii.

The posterior border of the skull roof is clearly marked by a ridge which, crossing
the flat table, passes outward and downward at the hinder end of the ornamented
surface of the squamosal and quadrat ojugal The smooth hinder surfaces of these
bones pass inward and backward round a concave cylindrical surface to end in a free

THE BRACHYOPID L AB YRINTHODONTS

335

margin, attached to a ridge on the hinder surface of the quadrate bone and its
cartilaginous dorsal extension. This ridge is supported on its admesial side by the
hinder edge of the pterygoid.

The ornament of the skull roof is of normal labyrinthodont pattern consisting of
pits — usually round and separated by low ridges with rounded upper surfaces — but
in some regions elongated into short grooves. It is an interesting confirmation of
Bystrow's view that the areas with grooved ornament represent regions of relatively
large growth in the direction of the grooves, that the grooves on the skull of B. watsoni
seldom run antero-posteriorly, but often transversely. They are best marked where

P.Tcm.Fos.

TEXT-FIG. 8. — Batmchosuchus bvowni Broom. Drawn (with some of the bones put back
into place) from a cast of the type skull in Cape Town, x \. I.Fr., interfrontal ;
P.Tem.Fos., post- temporal fossa.

they run into the suture between the supratemporals and the squamosals — implying
a widening of the skull here — and on the postorbital at its suture with the squamosal,
which in continuation of them has very small round pits, suggesting that in youth the
postorbital was wedged in between the squamosal and supratemporal. Through
this normal ornament the lateral line grooves make their way as wide but shallow
troughs, usually very broadly V-shaped in section. Their flanks often bear close set,
low, transverse ridges. They are very extensively developed, with a normal
distribution, though there is a possible transverse groove on the left tabular to which
it is difficult to find a parallel.

336 THE BRACHYOPID LABYRINTHODONTS

The occiput was correctly shown in my figure 28 (1919), but further preparation
has been made of the left stapes. This is now shown to rise more nearly to the
under-surface of the tabular at the admesial end of its upper border, so that the depth
of the bone is increased above the notch for a blood vessel, and the ventral surface of
the inner end is widened and its anterior part produced downward as a flange which
presumably had a sutural attachment to the outer border of the parasphenoid.
The pterygoid is shown to have a parotic plate reaching upward toward the cranial
roof, on the inner surface of the quadrate from the upper side of the horizontal shelf
which projects into the exoccipital and parasphenoid.

BATRACHOSUCHUS BROWNI Broom

This species is still represented only by the type specimen, which, when Broom
described it, was a split slab (? nodule), one side of which showed the visceral aspect of
the roof of the skull, the other had been prepared by him to show the palate. Sub-
sequently the South African Museum replaced the skull roof and prepared the orna-
mented outer surface, which, though a little broken and misplaced, is beautifully
preserved ; only the quadratojugals and anterior part of the face are missing. Its
structure is shown in Text-fig. 8.

It differs from B. watsoni in being proportionately longer and narrower, with the
eyes further back, and in the greater projection of the occipital condyles behind the
occipital border of the skull roof.

The lateral line grooves have the same general distribution as in B. watsoni, but
are actually and relatively much narrower and of more U-shaped section. There
is a peculiar inwardly and backwardly directed branch of the supratemporal line
on the hinder part of the postorbital, absent in B. watsoni.

The shapes of the individual bones of the skull roof are essentially the same,
though B. browni has an interfrontal bone between the frontal and nasals.

? BATRACHOSUCHUS sp.

In 1911 I collected three fragments of labyrinthodont jaw (D.M.S.W., B.I26) and
an atlas (D.M.S.W., 6.140) from a small weathered slope of shale of the Cynognathus
zone by the roadside on the farm Luiper Kop between Burghersdorp and Knapdaar
Two of the jaw fragments fit together to make the very weathered posterior end of a
right mandible, the other is part of the left ramus (including the front half of the
articulation) evidently of the same individual ; right and left sides supplement one
another and show the structure well.

The essential features (Text-fig. 9) are that the jaw is low, nearly as wide as high
at the articulation, with a well formed articular face behind which a long process
stretches backward. This has a core of articular, forming a median strip of its upper
surface, which is bordered by surangular and prearticular, the angular sheathing a
strip of its lower and outer surface. The details of structure differ greatly from those
of Capitosaurs, Benthosuchus, and the Spitsbergen Trematosaurs described by Tage
Nilsson, and there can be no real doubt that the jaw (which much resembles those of

THE BRACHYOPID LAB YRINTHODONTS

337

Bothriceps, Pelorocephalus and Dvinosaurus, and is comparable with that of Eobrachy-
ops) is that of a Brachyopid. If it be then presumably it belongs to Batrachosuchus,
which it fits quite well.

Remains of amphibia are so rare in the Cynognathus zone that " association "
of labyrinthodont fragments found close together is usually justified, and I therefore

TEXT-FIG. 9. — ? Batrachosuchus. (D.M.S.W., B.I26). x -£. Posterior part of a lower
jaw, a little restored from fragments of each side. A, Right ramus, outer surface ; B,
left ramus, inner surface ; c, right ramus, ventral surface ; D, right ramus, dorsal
surface. Ang., angular ; Art., articular ; Pr.Art., prearticular ; Sur.Ang., surangular.

describe the atlas found with these jaw fragments as that of Batrachosuchus^ The
atlas (Text-fig. 10), partly covered with a thin haematite film, is much weathered,
not completely prepared, and has lost its neural arch. But it is a single bone, only
about half as high as it is wide with a laterally concave posterior surface, widening a
little anteriorly. The anterior face bears two shallowly spheroidal hollows for
articulation with exoccipital condyles which are separated by a rather wide, slightly

338

THE BRACHYOPID L ABYRINTHODONTS

hollowed and pitted surface. The floor of the cavity for the spinal cord is wide and
somewhat depressed over its length ; the roots of the neural arch are, so far as can be
judged, extremely slender. The lower surface of the bone is pitted by irregular,
often longitudinal depressions. The specimen measures 4-8 cm. across the condylar
depressions. The condyles in B. browni are 6-0 cm. across. In B. watsoni the
corresponding width is 5-0 cm., only a few millimetres greater than that of the atlas,
and the same depth. The only important recognizable feature is the absence of any
indication that the atlas was ever more than a single bone.

TEXT-FIG. 10. — ? Batrachosuchus. (D.M.S.W., 3.140). x f . Atlas associated with the
original of Text-fig. 9. A, From below ; B, from in front ; c, from above ; D, from
behind.

PELOROCEPHALUS MENDOZENSIS Cabrera

The specimen, a skeleton, was found as a weathered impression to which films of
bone adhered, showing the palate and misplaced lower jaw. The palate, as Cabrera
rightly recognized, is extremely like that of Batrachosuchus, agreeing in general
proportions very closely with B. browni though (very probably owing to the imperfect
preservation) the subtemporal fossa is rather wide. The condyles agree, and the
suture bounding the parasphenoid is shown sufficiently to make it evident that the
exoccipital met the pterygoid in a long suture. The lower jaw shows an anterior
border to the articular facet exactly similar to that in the jaw referred above to
Batrackosvckus, and the remains of a similar long retroarticular process. The
specimen is preserved in Buenos Aires.

" PLATYCEPS " WILKINSONI Stephens

The following account is based on an enlarged photograph of the type specimen,
which I actually handled about 30 years ago, (PL 39 and Text-fig, n). The specimen,

THE BRACHYOPID LABYRINTHODONTS

339

which extends backward to show some 25 body segments, is seen from above as dark
grey crushed bone on a grey-yellow, fine-grained sandy matrix. The lower jaws are
in position and the head, shown from above, is widened by being flattened almost into
a plane. The skull is widest across the quadrate condyles and is semi-elliptical.
The occipital border is straight and lies behind the quadrates, there being some
rounding of the hinder edge of the tabular to enable it to pass directly into the hinder
border of the ornamented outer surface of the squamosal. Squamosal and
quadratojugal pass inward, behind the quadrate, and then backward. The jugal

Tab.

RC.+. Proc.

I.CIav.

TEXT-FIG, n. — " Platyceps " wilkinsoni Stephens, x if. Tracing of the skull, branchial
arch skeleton and dermal shoulder girdle, made from an enlarged photograph of a speci-
men (the type) from Gosford, New South Wales, now in Sydney. (Compare with
Plate 39). Br.I-IV, branchial arch skeletons I -IV ; Clav., clavicle; D.S.O.,
dermosupraoccipital ; Hy., hyoid arch skeleton ; I.CIav., interclavicle ; Ret. Proc.,
retroarticular process of the lower jaw ; Sq., squamosal ; Tab., tabular occipital process.

is extremely shallow lateral to the large orbit, and the shallow maxilla stretches
backward below it. The orbits are widely separated, of considerable size, but leave a
relatively large preorbital face. The tabulars and dermosupraoccipitals bear powerful
occipital flanges which pass backward and inward, showing that the condyles
must have been placed far back, though it is evident that the dermal processes
have been flattened out into the plane of the skull top and the extent of their
projection back thus exaggerated. The retroarticular part of each jaw ramus
is visible as a long projection.

340 THE BRACHYOPID LABYRINTHODONTS

The most striking quality is, however, the presence of long bony ceratobranchials.
The ceratohyal is, perhaps, visible on each side of the head as an obscure stain mesial
to the long retroarticular process, but four ceratobranchials are clearly shown on each
side between this element and the outline of the body as suggested by the clavicle.
Each ceratobranchial is a bony rod lying close pressed against its fellows, the anterior
ends of the rods seem to lie along a line placed obliquely, its outer end further for-
ward than its mesial point. The chief bones in all the arches are of much the same
length, but it is evident on the left side that the first ceratobranchial is greatly ex-
tended by some structure — presumably cartilage — for a very long distance, whilst on
the right side the last ceratobranchial has three or even more independent rings of
bone surrounding its distal end, and there is a suggestion of a very extensive spread
of soft structures beyond them.

The vertebral column is represented by paired series of structureless bony masses.
These seem to be in general paired ossifications in the neural arches, with slight
suggestions of other paired ventral bones. The ribs are short, nearly straight bones,
widening proximally in a manner which suggests that they may have had double
cartilaginous heads, but actually ending as bone some distance lateral of the vertebrae.
The distal ends of the ribs sometimes widen, possibly as a result of crushing of a very
thin bony cylinder. The first three vertebrae seem to be somewhat more massively
ossified than those further back, but curiously there seem to be no ossifications in the
exoccipital condyles.

The dermal shoulder girdle is well shown from above, the structure of its lower
surface " printing through ". The interclavicle is large, two-thirds as long as the
skull table, and its anterior end seems to have been almost semicircular, the widest
point being about at mid length. From here its lateral borders pass inward and
backward to end abruptly at a nearly straight transverse posterior border. The
clavicles have a long overlap on the ventral surface of the antero-lateral parts of
the interclavicle, and probably just meet at its anterior end. Their ventral parts are
expanded, the dorsal process apparently turning a little backward and being capped
by a cleithrum.

The scanty ossification of the vertebral column and the apparent lack of ossification
in the exoccipitals suggest that Platyceps is very young, as indeed its very small size
in comparison with other Brachyopids makes probable. The well ossified cerato-
branchia thus imply that the animal retained its external gills into adult life, and was
neotenous, as seems to have been the case in the " Plagiosaurid " Gerrothorax, where
Nilsson (1945) has recorded the existence of ossified ceratobranchials in an obviously
adult animal.

" PLAGIOSAURIDS "
DESCRIPTION OF MATERIAL

The genus Plagiosternum, established by E. Fraas (1889) for fragments from the
Upper Muschelkalk of Crailsheim, has had other specimens referred to it. But the
skull is still represented by fragments whose areas do not overlap, so that all that can
be said about it is that it may well have resembled those of Plagiosaurus and Gerro-

THE BRACHYOPID LABYRINTHODONTS 341

thorax. The skull described by Fraas (1913) as " Plagiosternum " pulcherrimum (now
referred to the genus Gerrothorax) is the only one belonging to the group which is
undistorted and really well preserved, but unfortunately the hinder surface of the
quadrat ojugal and quadrate is broken off on the right side and completely missing on
the left, and sutures are difficult to trace on the very richly ornamented outer surface.
Von Huene (1922) was the first to attempt to do so. Tage Nilsson (1937) has
produced another version which for the postorbital part agrees reasonably with von
Huene's paper, but differs absolutely for the more anterior regions. Nilsson's
account of a Swedish specimen, Gerrothorax rhaeticus, includes a description of a strip
of the hinder part of the dorsal surface of a skull whose dorso-ventrally compressed
occiput is also shown, and inspection of the material shows that his excellent figures
make the structure evident. Thus we have evidence in support of Nilsson's reading
of the difficult skull now in Stuttgart.

A fragment of the right hinder corner of a skull from Halberstadt, the type of
Jaekel's Plagiosaurus depressus, is illustrated by Nilsson in an excellent photograph,
and the structure of its dorsal surface is evident. It is differentiated from all others
by the meeting of the dermosupraoccipital and postfrontal (!) so that the parietal
does not touch the supratemporal.

The only other described genus is Plagiosuchus von Huene (1922). This is
represented by a skull whose palate is largely shown, two misplaced lower jaws, an
atlas, some ribs, a scapulocoracoid and clavicles and interclavicle, in articulation
but incomplete posteriorly. There are no ways in which this specimen can be
compared in detail with other Plagiosaurs, but it is important because its lower jaw
has an immense retroarticular process, similar to, though proportionally longer than
that which I have described above as Batrachosuchus. There is also an atlas, a single
bone again much like the one I have referred to Batrachosuchus.

POINTS OF AGREEMENT WITH OLDER BRACHYOPIDS

Thus the Plagiosaur skull is reasonably well known : it agrees with that of the
older Brachyopids in such important characters as :

(1) The way in which, as shown in Gerrothorax rhaeticus, G. pulcherrimus and
Plagiosaurus depressus, the pretympanic flange of the squamosal wraps round the
outer side of the quadrate and forms a laterally concave face on the occipital surface,
ending, as far as one can judge from the somewhat damaged material, in a ridge
which faces a similar border of the pterygoid, the nature of the quadrate not being
clearly shown.

(2) The up-turning of the lateral wings of the pterygoids from the subtemporal
fossae, so that the palate forms a broad D-shaped arch (seen in G. pulcherrimus,
less well in G. rhaeticus and Plagiosternum granulosum) .

(3) The fact that the occipital condyles lie far behind the dermosupraoccipital.

(4) The absence of an otic notch.

(5) The broad parabolic outline.

(6) The existence of a very large retroarticular process of the lower jaw.

342

THE BRACHYOPID L AB YRINTHODONTS

(7) The extreme shallowness of the anterior part of the skull to the hinder end of
the maxilla, and the deepening seen in side view, from a point behind the orbit to
the quadrate condyle (cf. Text-fig. 12).

The differences in the pattern of the dermal bones of the skull roof, which distinguish
the various genera, are obviously due to " improvisations " to enable a series of
bones growing outward from centres more or less fixed to fill a steadily contracting
area. Even in Batrachosuchus this difficulty is seen in the preorbital surface ; in the
Plagiosaurs it spreads to the temporal region.

Thus the close relationship between the Upper Permian and Lower Triassic
typical Brachyopids and the Middle and Upper Triassic Plagiosaurs is obvious,
depending on special detailed common peculiarities not known in any other con-
temporary Labyrinthodonts.

Orbif

H.

Tab.

P. MX.

Ex.Oc
Cond.

TEXT-FIG. 12. — Gerrothorax pulcherrimus (Fraas). Drawn from a cast in the Museum of
Zoology, Cambridge, x f approx. Lateral view of the skull based on a photograph
taken at a distance of some 12 ft., and then enlarged : it is thus very nearly a parallel
projection. Ec.Pt., ectopterygoid; Ex.Oc.Cond., exoccipital condyle ; MX., maxilla ;
P. MX., premaxilla ; Pt., pterygoid ; Qu., quadrate ; Qu.J., quadratojugal ; Sq.,
squamosal ; Sub.Tem.Fos., arrow directed into the subtemporal fossa ; Tab., tabular ;
X, tenth cranial nerve foramen.

DVINOSAURUS COMPARED WITH OLDER BRACHYOPIDS

In 1919 I suggested that the then undescribed Dvinosaurus from the Russian
Upper Permian was a possible ancestor of the Brachyopids, pointing out (1926 :
199) the existence of bony branchial arches and suggesting that the animal was
neotenous. In 1936 a description of Dvinosaurus by P. P. Sushkin was published
posthumously ; this is still of great value, even though in 1938 Bystrow gave us a
beautifully illustrated account and discussion of the animal. Sushkin accepted the
view that Dvinosaurus and Batrachosuchus are close relatives, though demurring —
obviously rightly — that, as Dvinosaurus is neotenous, it cannot well be an actual
ancestor of the other form. He was at that time the only man who had examined
both animals, and he added to the set of resemblances I had set out the remarkable
similarity between the stapes of the two animals, a matter of importance, for the
massive bone differs greatly from those of all other Labyrinthodonts.

THE BRACHYOPID LAB YRINTHODONTS 343

Bystrow agrees that in general appearance the skull of Dvinosaurus resembles the
Brachyopids, but finds (pp. 271, 272) a series of differences which are :

(1) Batrachosuchus, though very large, lacks ossified basioccipital and supra-
occipital.

(2) In Batrachosuchus the parasphenoid and the pterygoids are united by a strongly
toothed suture, whereas in Dvinosaurus this arrangement is lacking. If there be an
attachment between these bones in Dvinosaurus it has no resemblance to the long
" sutura paraspheno-pterygoidea " of Batrachosuchus,

(3) Other matters in which Batrachosuchus differs still more from Dvinosaurus are
the reduction of the paroccipital, the loss of any trace of the intertemporal, the
proportionately short palatal ramus of the pterygoid, and the passage of the sulcus
jugalis (of the lateral line) along the squamoso-quadratojugal suture.

Most of these differences are the results of evolutionary change of characteristically
labyrinthodont type, since : —

(1) All early (Lower Permian) Labyrinthodonts have a well-developed basioccipital,
some few a supraoccipital, bone. All Upper Triassic Labyrinthodonts lack both
bones.

(2) Several Lower Permian, and nearly all Upper Carboniferous, Labyrinthodonts
have a movable articulation of the pterygoid on the basipterygoid process. All
Triassic Labyrinthodonts have a long sutural attachment of the pterygoid to the
parasphenoid.

(3) All Lower Permian Labyrinthodonts (except Eobrachyops) have an ossified
paroccipital — in part visible from behind — but no Upper Triassic form retains that
condition, the paroccipital ceasing to be ossified and being concealed from view by
a meeting of the exoccipital and tabular. An intertemporal bone exists in Seymouria-
morphs of all ages, and in many early Labyrinthodonts — for example, it is present in
Loxomma but has been lost in all other Loxommids. Amongst Rhachitomi an inter-
temporal is present in the so-called Edopsids (most of which are of Pennsylvanian age)
and in Romer's Trimerorhachoidea (mostly Carboniferous, with some early Permian
descendants). All other Rhachitomi and Stereospondyli lack an intertemporal,
obviously by loss. Thus its occurrence in Dvinosaurus is merely the retention of a
primitive character.

By a comparison between all then-known Lower Permian and all Triassic Laby-
rinthodonts I showed in 1919 that a great enlargement of the interpterygoid vacuities
was a characteristic change found in that group as it is traced from its beginning to
its end. Thus the reduction in the length of the palatal ramus of the pterygoid
in Batrachosuchus as compared with Dvinosaurus is exactly what would be
expected.

The passage of the sulcus jugalis along the squamoso-quadratojugal suture in
Batrachosuchus does not exist ; Bystrow was unfortunately misled by my published
photograph.

The only other difference is that in Dvinosaurus the interclavicle has a long narrow
posterior extension and a median incision as a long narrow slit anteriorly, whilst in
" Platyceps " the interclavicle has a wide posterior end terminating in a gently rounded
but essentially transverse border, and no indications of an anterior median slit. The

344 THE BRACHYOPID LABYRINTHODONTS

clavicles of Dvinosaurus nearly approach one another but are narrow, whilst those
of " Platyceps ", also not in contact with one another, are greatly widened ventrally.
There is thus a real difference between Dvinosaurus and the later forms.

But Dvinosaurus is neotenic : it retains a larval condition physiologically and
structurally. It is true that " Platyceps " is larval and probably neotenic, as is Gerro-
thorax. But it is probable that the extent to which larval qualities were lost in
neotenic Labyrinthodonts could vary, for metamorphosis was obviously a long
process in Labyrinthodonts, for example in Archegosaurus , where an animal with a
skull length of 7-0 cm. may still retain the denticles guarding the inner openings of
the branchial clefts. Narrow lower ends to the clavicles are found in Eryops and in
Branchiosaurs, and may well be a persistent larval feature.

Romer (1947 : 125) accepts Bystrow's view, and elaborates a comparison with
Trimerorhachis and Saurerpeton. But Sushkin had, with a true instinct, recognized
that in Dvinosaurus ..." there is a well differentiated processus internus of the
pterygoid, tapering admesially ; its posterior edge being thickened and bearing a
facet for the articulation with the parasphenoid ; the articular surfaces of the
pterygoid and parasphenoid fit completely and there is no ground for supposing that
there existed also a junction of the pterygoid with a true processus of the chondro-
cranium which may not have been preserved in the fossil condition ". He recognized
a real resemblance to Trimerorhachis in this matter. It is evident that the condition
is unusual, the original basipterygoid process having vanished in association with a
great thinning of the basisphenoid, a new articulation between membrane bones
having taken its place. I can see no reason why the parasphenoid and pterygoid so
brought into apposition should not later gain a sutural connection, as they did in the
development of Eryops from an Edops-like ancestor where, however, the basisphenoid
core of the process remains (even long after the sutural attachment of the two
membrane bones has spread into a long flat sheet) as the cartilaginous infilling of the
" conical recess ". Thus this character is no evidence against a close association
between Dvinosaurus and the Brachyopids.

I would add to the features in which Dvinosaurus resembles the Brachyopids the
fact, shown in the two side views of the skull and lower jaw published by Sushkin
(1936 : 60, fig. 4) and Watson (1951 : 102, fig. 42), that the face is very shallow, the
temporal region relatively deep.

EOBRACHYOPS TOWNENDAE gen. et sp. nov.

DESCRIPTION OF MATERIAL AND MANNER OF FOSSILIZATION

Some twenty-five years ago I found, in the mass of unattractive, unregistered
Texan Permian material belonging to the Cope collection in the American Museum
of Natural History, a small Labyrinthodont skull almost entirely covered by a most
unpromising matrix. This was mainly a very hard, dark red, cemented mudstone,
partly represented by spheroidal nodules and for the rest by the light green matrix
in which bone structure is usually destroyed. This specimen I prepared with a

THE BRACHYOPID LAB YRINTHODONTS 345

hammer and needle-pointed chisels in odd hours during the early part of the late war
when I was fully engaged in government work. The preparation is satisfactory,
such matters as the surface ornament and sutures on the outer surface being perfectly
shown. But the specimen has been damaged, and it is necessary to explain its
condition in some detail to justify the restored drawings which I now publish, and
to bring out its eventful history, which has a bearing on the mode of deposit of the
rocks in which it was buried.

The skull, very shallow in front, is deep in the occipital region. The extreme end
of the nose, and part of the anterior surroundings of the left orbit, had been broken
off and lost before collection, and the right quadrate region is weathered away. The
animal after death was disturbed ; the left squamosal turned outward into the plane
of the upper surface, the left pterygoid disarticulated and carried over to the right
side of the palate. It now rests on the ventral surface of the right clavicle, which has
moved forward so that its dorsal ramus passes up through the right interpterygoid
vacuity to the roof of the skull. The left ramus of the lower jaw is misplaced so that
its inner surface faces upward, lying below, and separated by some 5-10 mm. from,
the visceral surface of the left squamosal. The anterior part of the jaw was not
collected. Below this jaw lies the interclavicle with its anterior end forward. On
the ventral side of this bone lie a number of vertebral elements, whilst the ventral
surface of the right clavicle is underlain by some ribs and a number of dermal scales,
of which others are to be found elsewhere. Such disturbance seems to me of a kind
which might have been brought about by pulls given by small animals at a time
when a good deal of the skin and other soft parts were still present, and the
whole appearance is more consistent with a drying carcass than with one under
water.

After these displacements the skull was fractured, two cracks crossing the table
from side to side, the central strip so separated being depressed 2-3 mm. below its
surroundings. At the same time the upper end of the dorsal ramus of the clavicle
was forced through a parietal, the fragments so formed standing up round its tip
like a tent, and the exoccipitals were fractured and forced up relative to the descend-
ing flanges of the dermosupraoccipitals. It is obvious that these fractures can only
have been made by pressure exerted by a heavy soft body into which the tent of
parietal fragments, still held together by skin, could have been forced. And the
only such structure I can think of is the foot of a large animal : and an animal
foot could only exert sufficient pressure in air.

Despite this damage it is possible to make reconstructions which must be nearly
accurate, as the right squamosal is in its natural position, though the quadratojugal
has been forced upward within it and has lost its lower margin. The orbital part of
the skull is extremely flattened in the actual specimen, but seems to retain very nearly
its original shape. The extreme depth of the cheek at the quadrate was suggested
by continuing the anterior part of the border of the quadratojugal backward, and
estimating the place of origin of the surface ornament of that bone from the radial
arrangement of its surviving parts. The point so determined is confirmed because it
enables a drawing of the lower jaw (previously made in what appeared to be a natural
position) to have an obviously natural occlusion when placed in articulation with the

346

THE BRACHYOPID LAB YRINTHODONTS

inferred quadrate condyle. The reconstructions were made by Miss J. Townend
after we had agreed on all matters of detail.

SKULL ROOF

The remarkable features of the skull roof are the small size, wide separation, and
very anterior position of the orbits, and the absence of an otic notch. All the bones
of a normal Labyrinthodont skull roof are present, those which build up the " table "
covering a very unusually large part of the whole area.

MX

No.

Par

S.Tem

SH.

D.S.Oc.

Qu.

Ex.Oc.

S.Oc.

TEXT-FIG. 13. — Eobrachyops toivnendae gen. et sp. nov. Reconstruction of the type
specimen (A.M.N.H. 2455). x i|. Dorsal aspect of the skull, viewed at right angles
to its upper surface. D.S.Oc., dermosupraoccipital ; Ex.Oc., exoccipital ; Fr., frontal ;
I. Tern., intertemporal ; Ju., jugal ; Lac., lachrymal ; MX., maxilla ; Na., nasal ;
P.O., postorbital ; Pal., palatine ; Par., parietal ; Pr.Fr., prefrontal ; Pt., pterygoid ;
Pt.Fr., postfrontal ; Qu., quadrate ; Qu.J., quadratojugal ; S.Mx., septomaxilla ;
S.Oc., supraoccipital ; S.Tem., supra tern poral ; Sq., squamosal ; Tab., tabular; X,
tenth cranial nerve foramen.

THE BRACHYOPID LAB YRINTHODONTS

347

The large parietals surround a transversely widened pineal foramen, meet the
frontals anteriorly, and are bordered laterally by the postfrontals, intertemporals and
supratemporals. Posteriorly they are attached to the dermosupraoccipitals which
stretch outwards behind the supratemporals to sutures with the small tabulars. The
presence and relations of the intertemporal are quite certainly shown ; it lies between
the parietal, postfrontal, postorbital and supratemporal — the position it holds in all
those Labyrinthodonts in which its occurrence has been described.

The squamosal is a remarkable bone, standing nearly vertically on the side of the
skull, articulating above by a long straight suture with the supratemporal and tabular
with its anterior end wedged in between the long postorbital and the jugal, with
which it has a very long suture. The upper part of the hinder border of the bone is
narrow and is separated by a now empty space from the upper part of the posterior

STam.

I.Tem.

P.O.

PhFr.

Pr. Fr.

D^n.

S.Mx.

P. MX.

TEXT-FIG. 14. — Eobrachyopstownendae. X ij. Lateral aspect of the skull and lower jaw.
Reference letters as before with : B.Oc., basioccipital ; Den., dentary ; P. MX., pre-
maxilla.

margin of the quadrate ramus of the pterygoid. Ventrally the hinder edge of the
squamosal is turned (over a rounded surface) abruptly inward nearly at right angles
to its outer surface and extends inwards toward, though it does not come into contact
with, the lower part of the hinder border of the pterygoid. The inner border of this
part of the squamosal is turned backward, no doubt applied to a narrow ridge of the
cartilaginous quadrate whose other surface was coated by the pterygoid.

The arrangement of the bones surrounding the orbit is shown with certainty.
There is a large postfrontal, articulating with the postorbital, intertemporal, parietal
and frontal bones, which ends anteriorly at a suture with the prefrontal. The
prefrontal is attached by its inner border to the frontal and nasal, and most unusually
extends forward to form part of the border of the nostril. The posterior part of the

GEOL. II, 8. 30

348

THE BRACHYOPID LAB YRINTHODONTS

lateral border is in contact with the lachrymal, the remainder with a bone whose
nature has to be discussed. This bone is excluded from the orbital margin, its
posterior end being attached to that upstanding process of the lachrymal which
reaches the prefrontal. From here forward to the nostril the bone is held
between the prefrontal and the maxilla, its anterior border forming a good deal
of the margin of the nostril. Except for a very remote possibility that the lachrymal
may only be a broken anterior part of the jugal, the neighbours of this puzzling bone
are certainly determined, and the only homologue which can be found for it is
the septomaxilla, which sometimes has a superficial exposure.

The lachrymal is a small bone forming the anterior and lower corner of the orbital
border. It is a little misplaced, but a notch bounded by the prefrontal and septo-

Tab. D-S/0c- foe.

TEXT-FIG. 15. — Eobrachyops townendae. x i£. Occipital aspect of skull viewed parallel
to the dorsal surface. Reference letters as before with : P.Tem.Fos., post-temporal
fossa ; Par.Ot.Pt., parotic flange of pterygoid ; Par.Sp., parasphenoid ; X, foramen for
the tenth cranial nerve.

maxilla shows its original position, and the mode by which it articulates with the
upper surface of the maxilla is obvious. The bone turns inward to form a wide wall
to the orbit, and is apparently fused with the upper surface of the palatine bone.

The jugal forms the posterior and lower corner of the orbit, meeting the lachrymal
in a suture which has been pulled apart so that the depressed surface, with an abrupt
posterior ending which housed the hinder end of the lachrymal, is very clearly seen.
Behind the orbit the jugal continues backward, between the postorbital and squamosal
above, and the maxilla below, until it reaches the incompletely preserved quad-
ratojugal.

The quadratojugal has a long exposure on the lateral surface of the skull, then
turns in to continue downward the posterior surface of the squamosal.

THE BRACHYOPID LABYRINTHODONTS
OCCIPUT

349

One of the more remarkable features of this skull is the amount by which the occiput
extends behind the posterior border of the ornamented table. In consequence the
occipital flanges of the dermosupraoccipitals are largely visible from above as a wide

p. MX.

MX

Pal

Ec

Par.Sp.

TEXT -FIG. 16. — Eobrachyops townendae. x i£. The palate viewed at right angles to the
parasphenoid. Reference letters as before with : Ec.Pt., ectopterygoid ; Par.Sp.,
parasphenoid ; Proc.Int.Pt., processus internus of the pterygoid ; Pt.Fac., dorsally
placed process of the pterygoid perhaps for the epipterygoid ; Vo., vomer.

triangle with a truncated apex. These flanges are marked off by a transverse groove
lying below and behind the rounded ridge, which is the hinder border of the table.
Each flange is notched posteriorly by a groove which passes forward near the middle
line below the skull roof. The lateral borders of the flanges stand out freely above
the post-temporal fossae.

The tabular has a very slender occipital flange, not visible from above, which
stretches inward and downward behind the cartilaginous paroccipital.

35<> THE BRACHYOPID LAB YRINTHODONTS

In occipital view the basioccipital forms a large, concave condyle, the " articular "
surface being spheroidal, though notched on its upper surface for the notochord.
The lateral borders are rounded and thickened. The upper surface is largely covered
by the lower ends of the exoccipitals, which send inward flanges (which probably
did not meet) below the foramen magnum. The exoccipitals then rise as columns on
each side of the foramen magnum, approaching one another until they are broken
across and now show a damaged upper surface. Their apparent continuation over
the summit of the foramen magnum is by a small bone (or bones) which is notched
below, and extends down the sides of that opening so as to meet them with a con-
tinuous admesial surface, there being no trace of the step which in most Labyrintho-
donts receives a cartilaginous supraoccipital. It is therefore likely that the bone in
question is actually a supraoccipital. From its posterior surface each exoccipital
extends forward and outward, surrounding the vagal foramen, and is perforated
by two more ventrally placed small openings which (by comparison with Capitosaurus)
may be for the Xllth cranial nerve. The rest of the bone clearly sheathed the
posterior surface of the persistently cartilaginous paroccipital, whose position is
fixed by that of the occipital flange of the tabular, which laterally sheathed its
posterior face.

PALATE

The parasphenoid is a remarkably large and elaborate bone. Its hinder margin
reaches the rim of the single concave occipital condyle, and indeed appears to border
the whole lower half of this structure, surrounding and hiding the basioccipital in
ventral view. Laterally the parasphenoid ends in a suture with the exposed lateral
surface of the basioccipital, but further forward this suture rises to cut the surface to
which the ventral face of the exoccipital is attached. From this place the para-
sphenoid widens, its border passing outward and forward to help to form an immense
basipterygoid process. In this region the bone is very shallowly concave mid-
ventrally, but then rises a little so that the anterior face of the process becomes a
deep and apparently vertical, transversely placed articular surface, which is applied
to a corresponding facet on the pterygoid. Mesial of this articulation the border of
the parasphenoid passes inward round the interpterygoid vacuity to extend anteriorly
as the outer edge of a processus cultriformis. The bone is then concealed by matrix
until what appears to be the ventral surface of its anterior end is exposed at the
anterior end of the specimen. The parasphenoid behind the basipterygoid process
has a nearly flat central region bounded on each side by a depressed area for attach-
ment of recti capitis muscles over the lateral parts of the basioccipital.

The pterygoid bears a powerful " internal process " which passes inward from the
meeting point of the palatal and quadrate rami. The straight borders of this
process meet at a point directed mesially at the hindmost place on the border of the
interpterygoid vacuity. This process is obliquely truncated by an apparently flat
articular surface opposed to the corresponding face of the parasphenoid ; the two
are separated in the specimen by a film of matrix about one millimetre thick. The
front face of the articular process of the pterygoid is obviously continued outward
and forward as the lateral margin of the interpterygoid vacuity. Lateral to and

THE BRACHYOPID LAB YRINTHODONTS 351

immediately behind the articular face the lower surface of the pterygoid is excavated
by a smoothly rounded pit, which lies immediately lateral to the tip of the para-
sphenoid. The floor of this pit extends inward onto a thin shelf of bone separated,
in the specimen, from the parasphenoid by about 2 mm. of matrix. It is in every
way probable that this space was originally occupied by a cartilaginous basipterygoid
process of the basisphenoid. Laterally the pterygoid turns down to form the
border of the subtemporal fossa, and, with the much deeper surface of the quadrate
ramus, makes a sheath to the temporal muscles, and bounds the characteristic
U-shaped section of the palate.

Further back the short quadrate ramus of the pterygoid is abruptly produced
downward as a deep flat sheet of bone with nearly parallel anterior and posterior
edges. The posterior border, originally applied to a cartilaginous quadrate, lies along
the hinder and inner border of the squamosal, but is throughout separated from it by
a narrow space originally rilled by the quadrate. Above the level of the inwardly
directed shelf, which lies above the presumed cartilaginous basisphenoid, the parotic
plate of the pterygoid rises vertically to, or very nearly to, the skull roof. The
anterior end of this part of the ramus plunges into matrix.

The only other visible feature of the pterygoid is a small triangular shelf extending
forward into the back of the interpterygoid vacuity from the anterior face of the
articular process. This has a flat ventral surface, and lies high in the skull, presum-
ably about at the level of the upper surface of the cartilaginous basisphenoid in its
vicinity. It may have supported the foot of an epipterygoid.

No part of the palatal ramus of the pterygoid is exposed except for its anterior tip,
seen through the orbit, which is grooved for attachment to the ectopterygoid and
palatine. The ectopterygoid is only represented by a single large tusk, whose length
is completely preserved.

The palatine bone is well shown. Its lateral border is throughout in contact with
the inner surface of the maxilla, and its inner margin — forming part of the border
of the interpterygoid vacuity — is completely visible. At its contact with the
ectopterygoid the palatal exposure of the bone is extremely narrow. Anteriorly, at
the level of the forward end of the vacuity, the bone bears two very large teeth ;
the anterior of these is a mature tooth of circular section, the tip being lost, the pos-
terior was in process of growth, arising within a ring of bone standing out from the
palatine. The crown of this tooth lies misplaced nearly horizontally and is sharp-
pointed, of circular section, and longitudinally striated. Mesial of these teeth the
bone extends inward, with a narrow exposure between the nostril and the vacuity,
to reach and be overlapped by the vomer. Anteriorly to the teeth the palatine
stretches forward internally to the maxilla until it meets the vomer, excluding the
maxilla from the internal nostril.

The vomer is a large bone well shown on the right side of the specimen. Its
contacts with the palatine have just been described ; it forms part of the border of the
interpterygoid vacuity, and has a process which extends backward along the lower
surface of the parasphenoid. It bears one large tooth, of which only half the root
is preserved.

352 THE BRACHYOPID LAB YRINTHODONTS

The premaxillae have been broken away, but a small process of one of them, resting
on the upper surface of the vomer, can be seen through the right external nostril.

The maxilla is a very shallow bone deepened anteriorly where it is attached to the
septomaxilla. It ends anteriorly in an almost vertical suture with the premaxilla.
Its lower border turns a little down posteriorly. It bears a series of small close-
set teeth, apparently uniform and circular in section, sharp-pointed and unexpectedly
high at the hinder end of the row.

LATERAL LINE

Lateral line grooves are very poorly represented. There is a somewhat unusual
groove which nearly surrounds the orbit. It begins about at the suture between the
prefrontal and lachrymal, and passes over the pre- and postfrontals and the post-
orbital on to the jugal, along which it passes — apparently just touching the lachrymal
and then descending on to the maxilla. There is a possible short branch crossing the
postorbital towards the intertemporal. The rest of the skull and the lower jaw seem
to lack any trace of these structures.

LOWER JAW

The hinder part of the left lower jaw is preserved, but is largely concealed by
irremovable matrix. It is of normal Labyrinthodont structure, but possesses some
unusual features. There is every reason to believe that its lower border was straight ;
there is a relatively long retroarticular process, a marked coronoid process, and a
lightly ossified articular. The angular is the usual boat-shaped bone with a radiating
ornament of ridges, and the hinder end of what is presumably the postsplenial is
similarly ridged. Little but the upper border of the surangular is exposed. This
bone extends back along the outer surface of the retroarticular process, in front of
which its upper edge bears a rounded notch marking the position of the articular
surface. From the anterior end of this notch a process passes inward, which laterally
carries a small part of the articulation for the quadrate, whilst mesially it seems to
have sheathed the front of the partly cartilaginous articular. The upper border of
the bone outside the supra-meckelian fossa is thick and rounded, ending at a clearly
marked suture with that hinder end of the coronoid which forms a definite process
with a narrow but rounded summit. The outer surface of the surangular has no
ornament so far as can be seen, and the dentary overlaps it.

The dentary reaches the ornamented surface of the angular and the postsplenial.
Its outer surface is rounded from dorsal to ventral edges, and is perfectly smooth.
The narrow dentigerous border supports a single row of small, close-set teeth of
circular section. Anteriorly a smooth surface of the dentary lies above the coronoid
on the inner surface.

The coronoid is applied to the inner surface of the dentary, rising to its process,
where it is wedged into the surangular. It surrounds the anterior end of the supra-
meckelian fossa and stretches forward on the inner surface of the jaw as far as the
specimen extends. This surface bears a patch of three small hemispherical denticles,
but is otherwise smooth. Its lower border rests on the prearticular.

THE BRACHYOPID LAB YRINTHODONTS

353

The prearticular is represented by a section of its hinder end separated by a space
(which represents the cartilaginous articular) from the retroarticular part of the
surangular. Anteriorly its lower margin is overlapped by the postsplenial.

Den.

Pt.Spl.

Sur.Anej.

Pr.Arf.

Den.

Cor.

Pr.Art.

TEXT-FIG. 17. — Eobrachyops townendae. x ij. The hinder part of the lower jaw. A,
Right side, external aspect ; B, right side viewed somewhat from below ; c, left side,
from within; D, left side from above. A ng., angular ; Art., articular ; Cor., coronoid ;
Den., dentary ; Pr.Art., prearticular ; Pt.Spl., post splenial ; Sur.Ang., surangular..

The very much bowed upper edge of the lower jaw makes it difficult to determine
its relation to the skull. It seems, however, clear that the jaw was so placed that the
lower border was turned in, the flat inner surface facing partly upward. Such a
position brings the upper part of the outer surface of the dentary (below its tooth
bearing edge) vertical, as it should be, and places the little of the articular cavity that
can be seen in a reasonable position.

354

THE BRACHYOPID LAB YRINTHODONTS
VERTEBRAL COLUMN

The vertebral column is represented by a single neural arch and a number of
" central " bones. All these are fractured and incomplete, but, so far as they go,
are well preserved.

The neural arch seems not to have been co-ossified with its fellow of the other side ;
it has a considerable neural spine expanded fore and aft at its summit. This spine
arises from an elongated body, along the side of the neural canal, bearing narrow
pre- and postzygapophyses which have no special character. The lower border of

Clav.

TEXT-FIG. 18. — Eobrachyops townendae. x i£. A, Reconstruction of the dermal shoulder
girdle from below. Areas in which the ornament is represented are seen on one side
or the other, borders in broken line restored, dotted area has not been seen on either
side, being hidden by matrix or other bones (its shape has no significance) . B, Recon-
struction of two vertebrae from the right side ; c, various scales. I.C., intercentrum ;
N.A., neural arch ; P.C., pleurocentrum ; Clav., clavicle; /. Clav., inter clavicle.

the bone is broken and does not show the way in which it articulated with the central
elements. There is, however, a short, anteriorly placed transverse process.

There are about a dozen incomplete circum-notochordal bones, all so much alike
that I have been unable to distinguish intercentra from pleurocentra with certainty.
They clearly surrounded a large notochord of the order of 8 mm. in diameter, their
outer surface being composed of dense bone with a good surface which tends to be
turned a little outward anteriorly and posteriorly. The bone is thin, about i mm.

THE BRACHYOPID LAB YRINTHODONTS 355

thick, and the inner surface is slightly roughened and not covered with dense periosteal
bone. All the fragments which show a natural termination end in a rounded point
extending beyond the periosteal outer surface. Those bones which I regard as
intercentra narrow dorsally from a maximum of 4-5 mm. to some 2 mm. in a height
of 8-0 mm. In all cases the wide ventral end is a fracture, and it seems probable that
the bones were really large segments of a cylinder and were not paired. These bones
show a narrow but rather deep flattening on the posterior part of the upper end of
their outer surface, which is presumably a rib-facet.

One bone, which seems to be complete, is 8 mm. high and has a maximum width of
2-5 mm. It is seen from its inner surface and is presumably a pleurocentrum. In
Text-fig. 1 8 I have made a restoration of two vertebrae on the basis of the remains
described above, and it may be mentioned as a point in its favour that it fits the
occipital condyle.

SHOULDER GIRDLE

The interclavicle and right clavicle are nearly complete, but are in part
hidden by matrix. The area of the ventral surface of the interclavicle on which
ornament is shown in Text-fig. 18 is exhibited on one or other side of the specimen,
and its middle line is fixed by a marked median ridge on the visceral surface of its
anterior end. Occasional short lengths of genuine margin, or of the overlap surface
for the clavicle, fix its shape within narrow limits. The right clavicle, seen from its
ventral surface, is very nearly complete, but much of it is covered by matrix and
other bones.

Thus the restoration rests on adequate evidence, and it seems to be certain that
the clavicles must have met for a considerable distance in front of the interclavicle.
It will be noted that the total width fits neatly that of the skull. The height and
direction, though not the structure of the dorsal process of the clavicle, is known ;
it stands up at right angles to the ventral surface of the bone and is 22 mm. high,
leaving room in the body for a cleithrum of normal proportions.

SCALES

In a number of places groups of small, oval, ? bony scales may be seen (Text-fig.
18). Each has an ornament of low concentric ridges. They appear to have over-
lapped extensively, and are about 3 mm. across, but very thin.

POSSIBLE RELATIONSHIP TO ACHELOMA CASEI AND TRIMERORHACHIS

The only animal to which the Labyrinthodont described above has any similarity
is that described by Broili in 1913 as " Acheloma " casei. The two specimens
agree in that they have a truncated posterior border to the dermal skull, with
no trace of an otic notch. In both the orbits lie far forward, with a well marked
smooth border surrounding each, and are not exceptionally close together.
Broili's lateral view seems to show a tooth-bearing fragment of maxilla not far below

356 THE BRACHYOPID LABYRINTHODONTS

the orbit, and shows that the cheek was deep. Indeed the two skulls not only
resemble one another in general shape, but in the character of their ornament and
the general direction of the ridges on the cheek bones which form it. They may well
belong to the same genus.

But they are specifically different. The new skull has a large, transversely
widened pineal foramen, " Acheloma " casei a very small one. The table of the new
skull is broader than it is in Broili's specimen, and so is the interorbital space, whilst
the orbit of "Acheloma " casei is wider than that of the new form. The squamosal
in "Acheloma " casei appears to be deeper than that of my animal, and the new

TEXT-FIG. 19. — Eobrachyops casei (Broili). x f. Reconstruction of the skull drawn
from Broili's figures, left side copied from right. The outline of the cheek in B, hypo-
thetical, is based on a comparison of the ornament shown on it with that of E. townendae.
A, Dorsal surface ; B, right lateral surface. Sq., squamosal.

skull being only some twenty-five per cent larger than the old, it is impossible to
explain the visible differences in proportion as due to growth. I propose to
call the new specimen Eobrachyops townendae gen. et sp. nov. Eobrachyops
expresses my opinion of its systematic position, townendae is in honour of Miss Joyce
Townend, who made the drawings which illustrate this and all my other recent work.
"Acheloma " casei Broili becomes Eobrachyops casei.

Eobrachyops comes from an unknown locality and horizon but belongs to the Cope
collection, much of which came from the Wichita formation, though some is Clear
Fork. Professor A. S. Romer has examined the specimen of E. townendae and tells
me that the matrix is certainly Clear Fork, as is E. casei It now bears the number
Amer. Mus. 2455.

The only contemporary form which resembles it at all is Trimerorhachis.

THE BRACHYOPID L AB YRINTHODONTS 357

Trimerorhachis has been described by Cope, Broili, Case, Broom and Williston,
but still needs much further description. I therefore give a further account of the
basicranial and otic regions.

The two genera have in common : a skull which is depressed, at any rate anteriorly,
anteriorly placed orbits, an intertemporal bone in the table, a shallow (or absent)
otic notch, an occipital condyle, the greater part of which is made by the basioccipital
— so that it is deep and has a concave articular face — and a " movable " articulation
between the basipterygoid process of the parasphenoid and the pterygoid.

TRIMERORHACHIS

VARIATION OF SPECIMENS

The following account is founded on four specimens which Professor Romer was
good enough to send me (M.C.Z. 1169, 1975 A, B & C) ; and on B.M.N.H., R-576,
which reached the Museum in 1885, from the collection of Waldemar Kowalevski,
to whom it had been given by E. D. Cope before 1883 (as is recorded in a letter
dated 30.vii.i885 from Cope to Henry Woodward in the archives of the Geology
Department of the British Museum).

The skull of Trimerorhachis is generally believed to have been very much flattened,
as much so perhaps as in a late Triassic Stereospondyl. But it seems certain that
this flattening, in some cases at any rate, did not extend to the occiput, although it
did to the quadrate regions of the skull.

The occipital condyle is always single, and concave, but it varies a good deal in
shape in different specimens — perhaps from different horizons. In one specimen before
me it measures 2-3 cm. transversely, and is 1-75 cm. high. Another is 2-25 by 1-5
cm., the outline being convex throughout. Another is 2-1 by 1-6 cm. and has
concave lateral borders below the middle of its height, whilst R-576 is 2-1 cm. wide
and only 1-2 cm. high, the basioccipital part of the structure being greatly reduced.
These variations are obviously related to the general flattening which takes place
with time in Labyrinthodonts. Associated with them is a change in the height of
the exoccipital above its condylar portion. In the second of the above specimens the
height is 1-3 cm., in the third 1-2 cm., and in the fourth only 0-75 cm., in other words
the flattening is general.

One of the specimens mentioned above (M.C.Z. 1169) has the table of the skull
well preserved and visible from both upper and lower surfaces and from the back.
The dermosupraoccipital bears a very obvious occipital flange 1-05 cm. deep, and this
" flange " is merely the occipital exposure of a thick mass of bone whose lower
surface, which lies nearly horizontally, extends forward for nearly a centimetre and
obviously rested directly on the upper end of cartilaginous continuations of the
exoccipital, and the paroccipital. There is no evidence of any overlap of the dermo-
supraoccipital on the occipital surface of the exoccipital. Thus the minimum height
of the occiput is 1-6 (condyle) + 1-2 (exoccipital) -f 1-05 (postparietal flange)
= 3-85 cm. (assuming — as we justifiably may — that it was vertical) and the width
of the table between the otic notches directly measured is 6-3 cm., i.e. as i : 1-64.

358

THE BRACHYOPID LABYRINTHODONTS

Eryops in Sawin's figure is i : 2-43. In fact the occiput of Trimerorhachis is deeper
and narrower than that of Eryops and even of Edops.

BRAIN-CASE

The basioccipital is a wedge-shaped bone forming the lower part of the single
occipital condyle. It has a short, free ventral surface posteriorly which extends

Pit.

Gr.B.Pf.Proc.
Art. Fac.

Por.Sp.

Pai.Ar.

Fac.B.Oc.

B

Art. Fac,

Proc.CuIt.

Pai.Ar.

TEXT-FIG. 20. — Trimerorhachis sp. (M.C.Z., 1975). x 2. Posterior part of the para-
sphenoid, with co-ossified basisphenoid. A, From above ; B, from in front. Art. Fac.,
articular face on the parasphenoid for the processus inter nus of the pterygoid ; B.Sp.,
basisphenoid ; Fac.B.Oc., ridged face for attachment of the basioccipital ; Gr.B.Pt.Proc.,
groove for the cartilaginous basipterygoid process ; Pai.Ar., foramen for the palatine
artery ; Par.Sp., parasphenoid ; Pit., depression for the pituitary body ; Proc.CuIt.,
processus cultriformis of the parasphenoid cut across ; T.V., transverse vein.

THE BRACHYOPID L AB YRINTHODONTS 359

outward to smooth lateral surfaces, usually concave, which terminate dorsally at the
attachment of the exoccipital. The greater part of the ventral surface is ridged for
the overlap of the hinder part of the parasphenoid, which turns upward round the
bone to continue its concave lateral faces forward over a suture, so forming a deep
pocket, presumably for the attachment of a recti capitis muscle. The dorsal surface
of the basioccipital has a median groove for the notochord, lateral to which lie the
facets for the exoccipitals, whilst anteriorly the bone thins until it gradually fades
away into the upper surface of the parasphenoid.

The parasphenoid is directly comparable with that of Eryops or — perhaps more
easily — a primitive Neorhachitome. It varies a good deal in shape in different
individuals, but these changes merely overlie a fundamental uniformity. On
its upper surface the parasphenoid at the root of the processus cultriformis is co-
ossified with an extremely thin basisphenoid. No dividing line between the two
bones can be seen in section, but that the basisphenoid does indeed exist is shown by
surfaces belonging to it not covered by perichondral bone, but formerly continued by
cartilage. These exist posteriorly toward the basioccipital, and laterally at the base
of the grooves, lying on the upper surface of the articular process, and bounded
behind by the lateral ridge, which show the position of cartilaginous basipterygoid
processes. Consideration of the facet on the pterygoid for these processes suggests
that the cartilage was shallow, the functional articulation being between the
parasphenoid and the pterygoid.

The dorsal surface of the body of the basisphenoid has a shallow concavity bounded
by a transverse ridge behind, which agrees with a similar structure in Eryops, on
whose floor Sawin held that the recti muscles had their origin. Further forward on
the upper surface, at the root of the processus cultriformis, lies a depression for the
pituitary, as in Eryops. Behind the lateral ridge on the upper surface of the para-
sphenoid is a smooth depression, passing out laterally, which no doubt sheathed the
lower part of the prootic. In one specimen (but not in R-576) a small foramen
enters the bone from the admesial end of this hollow.

The exoccipital has a base which is closely attached to, and ultimately fuses with,
the upper surface of the basioccipital. This base is moulded on the upper surface of
the notochord, its posterior surface forming part of the great concave " condyle ".
It expands inward to floor the brain cavity and nearly (or quite) meets its fellow in
the middle line. The base extends forward and a little outward until it ends abruptly
at a face presumably lying in contact with the lower cartilaginous portion of the
paroccipital. Anteriorly the upper surface of the base forms the floor of the vagal
foramen, whose posterior wall is the massive ascending part of the exoccipital which
bounds the foramen magnum. It is important to note the complete absence in
Trimerorhachis of that perichondral extension of the exoccipital, which in Eryops
and all its successors overlaps the paroccipital to an ever-increasing extent. The
hinder surface of the exoccipital which rises to the dermosupraoccipital is essentially
flat, but it bears a small protuberence a little above the level of the floor of the foramen
magnum. This recalls the facet for attachment of a proatlas in many reptiles and
may have had this function, for processes for the attachment of such a bone exist on
the anterior faces of the atlantal neural arches (Cope & Matthew, 1915, pi. 8, fig. ib).

360 THE BRACHYOPID LAB YRINTHODONTS

The upper end of the exoccipital is abruptly truncated and no doubt continued by
cartilage.

The paroccipital is well shown in three of my specimens. It is a large bone with
the posterior surface covered by a smooth perichondral layer. The admesial part
of its posterior surface forms a stout pillar, ending below in a rounded and laterally

Par. Sp

Gr.B.Pt.
Proc.

-Sp. Art. Fac.

Par.Oc.

Fen.Ov

B.Oc.

Ex.Oc.

TEXT-FIG. 21. — Trimerorhachis sp. (B.M.N.H., 11.576). x i^. Posterior part of the
parasphenoid with hinder part of the brain-case articulated with it (compare with Text-
fig. 20). A, From above ; B, from the right side ; c, from below. Reference letters as
before with : B.Oc., basioccipital ; Ex.Oc., exoccipital ; Fen.Ov., fenestra ovalis ;
Par.Oc., paroccipital ; Pr.Ot., prootic.

THE BRACHYOPID LAB YRINTHODONTS 361

widened mass clearly originally continued by cartilage, which is applied to — but
extends laterally of — the front end of the base of the exoccipital. A groove on its
hinder surface passing forward and inward completes the vagal foramen, and the upper
end of the bone must touch the exoccipital and nearly reaches the skull roof. The
bone is continued laterally by a massive paroccipital process, which is thin dorso-
ventrally but wide from back to front. The outer end of this process is unossified,
but clearly extended laterally as cartilage to be received in a depression on the lower
surface of the tabular bounded laterally and posteriorly by an upstanding ridge.
This depression lies far in front of the corner of the tabular at the inner side of the
otic notch. Throughout the paroccipital is widely separated from the skull roof by a
large post-temporal fossa. The lower surface of the paroccipital process is only
shown in R-576, where it extends forward to meet the corresponding surface of the
prootic, and admesially ends abruptly in a border lying parallel with the middle line,
which is the upper margin of the bony fenestra ovalis.

The prootic is ossified in R-576. It is a bone much resembling the paroccipital, to
which it is closely applied, the two together forming a wide but shallow process, with
a wide groove running antero-posteriorly above it, which flares out on to the shallow
anterior face of the prootic. Ventrally the prootic has a base descending into a mass
of cartilage resting on the parasphenoid, and it forms the anterior part of the fenestra
ovalis.

The stapes lies nearly in position on each side of one specimen (M.C.Z. 1169), from
which Text-fig. 22 is drawn. In it the stapes is placed in position and is necessarily
foreshortened. The bone has an almost straight outer border, which turns in at its
lower end to the accessory process. The shaft of the bone is gently concave across
its lower (posterior) surface, rounding over on the outer side, the inner forming a
distinct edge. The inner border bows out from the bone a little in the middle of its
length, but cannot have come into contact with the otic capsule. Distally the bone
ends in a small rounded surface, whilst proximally the powerful accessory process
turns a little inward and ends in a surface, at about 45 degrees to the length of the
bone, which is itself thrown out into minor processes for an attaching ligament.
This process is separated by a groove, into which opens the foramen for the stapedial
artery, and a notch from the more powerful process which carries the footplate.
This process passes inward at an angle of about 130 degrees to the shaft, is massive,
and truncated by a slightly concave surface for attachment to the membrane closing
the fenestra. It seems evident that this ancillary process was connected to the outer
end of the lateral ridge of the parasphenoid, and that the distal end of the bone lay in
the tympanic membrane, the footplate being thus brought squarely into the obvious
position of the fenestra ovalis.

The pterygoid has a remarkable mode of articulation with the basis cranii. It is
evident that the important attachment was between a face directed backward, and
placed vertically on the posterior surface of a special inwardly directed process of the
pterygoid, and the front face of the parasphenoidal process. Each of these two
opposed facets is concave, so that it is evident that the joint — though firmly fixed—
was flexible owing to the presence of a ligamentous pad between the two bones.
The outer end of the posterior border of the articular process of the pterygoid swings

362

THE BRACHYOPID L AB YRINTHODONTS

round so as to limit a pit whose hinder wall is the beginning of an ascending rounded
ridge passing upward and backward at the root of the quadrate ramus. Little
space is left for a contact between the cartilaginous basipterygoid process and the

Tob.

D.S.OC.

St.

Ex.Oc.

TEXT-FIG. 22. — Trimerorhachis sp. Reconstruction of the posterior part of a skull made
from specimen M.C.Z. 1169. About natural size. A, From behind ; B, from below (to
be compared with Text-figs. 20 and 21). Reference letters as before with : D.S.Oc.,
dermosupraoccipital ; Proc.Int.Pt., processus internus of pterygoid ; Pt., pterygoid ;
Pt.P.Tym., post-tympanic ridge of the pterygoid ; Qu., quadrate ; Q.J., quadrato-
jugal ; Sq., squamosal ; St., stapes ; Tab., tabular.

pterygoid. Apart from the special articular process (not otherwise known in
described Texas Labyrinthodonts) the pterygoid generally resembles that of Eryops,
though it possesses the groove bounded by a ridge on the posterior surface of the

THE BRACHYOPID LAB YRINTHODONTS

363

quadrate ramus, originally recognized by Sushkin as the floor of the tympanic cavity
in Capitosaurus. The remainder of the structure of Trimerorhachis is well described
by Cope, Broom, Williston and Case.

COMPARISON OF EOBRACHYOPS WITH TRIMERORHACHIS

RESEMBLANCES

Eobrachyops and Trimerorhachis are not very different in size ; the type skull of
T. conangulus Cope is about two-thirds as long as E. townendae, whilst most specimens

A .""^x B

TEXT-FIG. 23. — Skulls of Eobrachyops and Trimerorhachis reduced to the same width
enabling them to be compared. A and B, Eobrachyops townendae, dorsal and palatal
views ; c and D, Trimerorhachis sp., from Cope's figures of the type skull of the genus,
Broom's figures, and Case's 1935 materials.

of T, insignis are twice the size. The two animals resemble one another in the general
outline of the skull, in the forward position of the orbits, and in their flat dorsal
surfaces. The otic notches are small in Trimerorhachis, and absent in Eobrachyops.
An intertemporal bone is present in each. The occipital condyle of each is a nearly
circular concave face, largely made by the basioccipital. In each the pterygoid

31

364 THE BRACHYOPID L AB YRINTHODONTS

articulates " movably " with the parasphenoid by transverse faces. In each a
pocket — apparently for the insertion of the recti capitis muscle — lies on the lateral
surface of the parasphenoid not very far behind the articulation with the pterygoid.
In each form the mandible is remarkable at this period in possessing a well-developed
retroarticular process.

The vertebral columns are similar in the lack of fusion of the two halves of the
neural arches in the anterior region, in the delicacy of the circum-notochordal
elements, and in the large size of the notochordal sheath. The dermal shoulder
girdles are similar in the presence of rhomboidal interclavicles, and in the wide
expansion of the ventral parts of the clavicles, a condition otherwise unknown in the
Texan Permian Labyrinthodonts — except in Archeria (= " Cricotus "). The two forms
possess identical bony? scales. There is therefore a number of real resemblances
between the two forms, and a further analysis is necessary.

DIFFERENCES

The resemblance in general skull outline is purely superficial. The small otic
notch of Trimerorhachis housed a tympanic membrane in which the stapes ended,
Eobrachyops must have entirely lacked such a structure, no doubt a secondary
condition associated with some change in the mechanism of hearing.

In Eobrachyops the quadrate lies far forward on the level of the occipital condyle,
in Trimerorhachis it is relatively far back, at least in some specimens. In Eobrachyops
the quadrate articulation is carried down by the deep cheek and lies far below the
floor of the basioccipital, whilst in Trimerorhachis the quadrate condyle seems to be
on the same level as the bottom of the occipital condyle.

In Eobrachyops the orbits are widely separate from one another and extend so far
laterally that the depth of the jugal and maxilla below them is extremely small.
In Trimerorhachis conditions are quite different, the face below the orbit being very
deep. In the detail of pattern of dermal bones in the circumorbital region the two
animals differ very greatly. In Trimerorhachis the lachrymal is a huge bone,
extending from the nostril backward below the orbit to meet not only the jugal, but
the postorbital. In Eobrachyops it forms scarcely more than one half of the outer
border of the orbit, and does not reach the nostril.

Thus the occurrence of an intertemporal in each form is merely the retention of a
primitive character, and there is no reason to suppose that the few resemblances
between the skull roofs have any phylogenetic significance ; they have been derived,
no doubt, from a common ancestral pattern by entirely different courses of change.

The basicranial and otic regions differ in the larger contribution which the exoc-
cipitals make to the condyle in Trimerorhachis, and in the fact that in that animal
there is no extension of the exoccipital over the paroccipital, whilst in Eobrachyops
there is to such an extent that the vagal foramen lies entirely in the exoccipital.
In Eobrachyops the paroccipital remains entirely cartilaginous, in Trimerorhachis
it is well ossified.

In Trimerorhachis there is a special process from the parasphenoid which bears the

THE BRACHYOPID LABYRINTHODONTS 365

articular face for the pterygoid, but in Eobrachyops this face is carried directly on
the front of the great lateral wing of the parasphenoid.

The palate of Eobrachyops has larger interpterygoid vacuities than that of Trimero-
rhachis, as shown by the fact that the pterygoid does not meet the vomer. The two
differ in the details of the dentition, and Eobrachyops is unique in that the vomer and
palatine meet laterally to the internal nostril.

The lower jaw of Eobrachyops differs markedly from that of Trimerorhachis in its
straight ventral border. The circumchordal elements of the vertebral column of
Eobrachyops differ from those of Trimerorhachis in the much more parallel sides of the
intercentra, which do not possess the typical rhachitomous wedge-shaped form of the
latter genus. In the shoulder girdle the clavicles meet in front of the interclavicle
in Eobrachyops, and not in Trimerorhachis.

Two STRIKING COMMON FEATURES

Thus it appears that the two genera are not very closely related : their resemblances
are dependent on the retention of primitive qualities and a somewhat similar skull
shape. However, the special process of the pterygoid which articulates with the para-
sphenoid is common to both, and is a very unusual feature ; and the retention of a
deep basioccipital as the greater part of the nearly circular condyle, whilst the basi-
sphenoid is so greatly dorso-ventrally depressed that the movable attachment of the
pterygoid to the basis cranii is essentially by means of a special facet of the
parasphenoid, is a remarkable point of resemblance between the two genera, as it
seems to be found in few other animals.

CHARACTERISTICS OF EOBRACHYOPS ARE THE SPECIAL QUALITIES

OF BRACHYOPIDS

The most striking special characteristics of Eobrachyops are (a) that the occipital
condyle lies considerably behind the posterior margin of the skull roof ; (b) that the
border of the pterygoid at the subtemporal fossa is suddenly carried upward so
that the palate in this region is a broad D-shaped arch ; (c) that the quadrate
condyle lies far below the ventral surface of the parasphenoid ; (d) that a distinct
space, formerly occupied by a cartilaginous ridge on the posterior surface of the
quadrate, separates the hinder border of the pterygoid from those of the squamosal
and quadrat ojugal ; (e) that the outer surface of the squamosal and quadrat ojugal
passes round on to the posterior face of the quadrate and there forms a laterally
concave, nearly vertical surface.

These five characters are essentially those which I listed in 1919 (pp. 47-48) as
special qualities of the Brachyopidae. Their occurrence in Eobrachyops and in no
other Labyrinthodont except the Brachyopids and Dvinosaurus seems to me strong
evidence for believing that Eobrachyops was the " prototype " of this group, in the
sense that it was a Lower Permian representative rather than literally ancestral to
any one of the later members.

366 THE BRACHYOPID LAB YRINTHODONTS

CONCLUSION ON RELATIONSHIP OF EOBRACHYOPS TO TRIMERORHACHIS

AND BRACHYOPIDS

It is evident that Eobrachyops more greatly resembles Dvinosaurus and the
Brachyopids than does Trimerorhachis. The latter lacks any sign of the deepened
cheek which exists in all Brachyopids, and to a less extent in Dvinosaurus. It has
often a considerable production of the quadrate condyle behind the occipital condyle,
in contrast to the condition in Brachyopids. In Trimerorhachis the occiput is
invisible from above (or nearly so) in contrast to the sloping occiput and very
posteriorly placed condyles of Brachyopids, Dvinosaurus and Eobrachyops. In
Trimerorhachis the pterygoid at the inner border of the subtemporal fossa is bowed
inward mesially to the inner surface of the jaw, and is not at all produced downward.
In Dvinosaurus this border is ... " strongly inclined ventralwards by its end
correspondingly to the general vaulting of the palatal surface " (Sushkin, 1936 : 56),
but it does leave a small part of the subtemporal fossa visible within the lower jaw.
The pterygoid of the Brachyopids differs from that of Trimerorhachis in ways which
carry the differences shown by Dvinosaurus to their logical conclusion, but Eobrachyops
has the Brachyopid condition in an extreme form.

In Trimerorhachis there is a functional otic notch, and a stapes of normal Laby-
rinthodont type ending in it. In Dvinosaurus and Batrachosuchus the stapes is
greatly deepened distally, its end being tied down by ligament to that posterior
surface of the ridge of the quadrate which separates the occipital part of the squamosal
from the pterygoid. It is indeed not easy to imagine intermediate stages between
the two conditions, whilst the resemblance between the stapes of Dvinosaurus and
that of Batrachosuchus is obvious (cf. Sushkin, 1936 : 59).

In Trimerorhachis the lachrymal is immensely enlarged and separates the orbit
widely from the maxillary border, in Dvinosaurus (Bystrow, 1938) it is less extensive
and the space between the orbit and the maxillary border smaller. In Brachyopids
it is still further reduced.

The comparison can be extended to many other features with the same conclusion :
that Trimerorhachis may well have some real relation to Eobrachyops, but that the
latter in all ways makes a better ancestor for Dvinosaurus and the Brachyopids.

EOBRACHYOPS COMPARED WITH PELION

Sushkin, when he recognized the remarkable processus internus of the pterygoid of
Dvinosaurus and showed its resemblance to that of Trimerorhachis, could only add as
a parallel Archegosaurus as I figured it (1919, fig. 2). But it also occurs in the Coal
Measure Saurerpeton and Erpetosaurus described by Romer (1930, 1947) and Steen
(1931). I therefore examined Steen's materials of these animals in the British
Museum. Dr. Steen had prepared them by removing the surviving remnants of
bone by hydrochloric acid, thus getting exquisitely perfect impressions from which
she could make squeezes in plasticene or dental wax only. But, as Mr. Donald Baird
of Harvard showed me, it is possible to make casts from such moulds with rubber
latex loaded with an opaque powder. Casts so made are incomparably better than

THE BRACHYOPID LABYRINTHODONTS 367

any at Dr. Steen's disposal, and enable me to add some details to her perfectly
accurate accounts of these animals.

IDENTIFICATION OF PELION LYELLI

Specimen B.M.N.H., R.2&74, was described as Pelion lyelli, though Dr. Steen
(1931: 874) notes that it differs materially from Romer's description of two skulls
referred to that species in 1930. In 1947 Romer (without resolving the differences
which Steen mentions) refers the three specimens in question to Hoodie's genus
Saurerpeton, founded on one of the specimens. Pelion lyelli has as type a partial
skeleton so covered by a film of matrix that no real knowledge of its skull structure
can be gained.

To Pelion Romer (1947) now refers a skull (described by Moodie as Erpetosaurus
tabulatus) for which he founded a genus Branchiosauravus in 1930. Romer (1930,
fig. 7) draws (but leaves unshaded) a bone projecting behind the table. This bone
in Hoodie's photograph (1916, pi. 25, 2) seems to lie below the skull roof and to have
a pair, less evident, but a good match. It is possible that these are the remains of an
occiput, stretching backward behind the occipital border of the table as that structure
must have done in Steen's Pelion. Thus it is probable that the two animals belong
to the same species, Hoodie's being older than Steen's, as it is half as large again.

RE-EXAMINATION OF PELION

Steen's specimen of Pelion lyelli may be described from the cast as follows. The
skull lies on its dorsal surface, both lower jaws being in position, articulated with the
quadrate condyles. On the spectator's right (animal's left) side the cheek is
completely flattened out into the plane of the table, the lower jaw is seen — dorso-
ventrally compressed — directly from below, but with the border of the pterygoid at
the sub-temporal fossa squeezed down into contact with its inner surface. This
crushing has disarticulated the pterygoid from the basis cranii, which, although
crushed down on to the skull roof, is still nearly symmetrically placed with respect
to the eyes. The right lower jaw is in articulation with the quadrate, but has become
detached at the symphysis and straightened, by pressure, lies so as to show the whole
of its outer surface. Its lower border has squeezed and broken the lower part of the
pterygoid where it forms the border of the sub-temporal fossa. The right maxilla,
jugal and quadrat ojugal lie in nearly natural articulation with each other and with
the quadrate and lower jaw, and are inturned so that their outer surface is shown.
Steen, in her accurate drawing (text-fig. 15), shows the large parasphenoid with a
rather definitely marked short blunt-ended basipterygoid process. The new cast
shows that this structure has its anterior edge drawn downward, so that there is a
large somewhat concave face directed forward to which a corresponding facet of the
pterygoid was attached. On the spectator's left the corresponding process is shown
with a special " processus internus " of the pterygoid in articulation with its flat
front face. On the spectator's right Steen's fig. 15 shows quite accurately a strange
hook-shaped mesial edge to the pterygoid, which in her restoration she attached to

368

THE BRACHYOPID LABYRINTHODONTS

the outer end of the parasphenoid process, but in the same figure she shows with
a dotted line a break in the inner border of the palatal process of the pterygoid. It
is evident from the new cast that this is the real processus internus, with a flat surface
directed backward for articulation with the anterior deepened face of the para-
sphenoid, the whole agreeing exactly with the structure seen on the animal's right.
The pterygoid extends outward from this articulation as a relatively enormous sheet
of bone somewhat cracked and crumpled but evidently continuous to a border,
which is pressed against the inner surface of the lower jaw for a very long distance, to

Ot.N.

Cor.

Sur.Ancj.

TEXT-FIG. 24. — Pelion lyelli Wyman. Reconstruction of a skull (B.M.N.H., R.26y4).
X 2*2. A, Dorsal ; B, palatal ; c, right lateral aspect. Reference letters as before.

the middle of the orbit in fact. The lateral strip of this bone bears an ornament of
ridges and grooves, deeply cut and of most unusual character. The sculptured
strip extends very far forward, but ends posteriorly a little in front of the level
of the basitemporal articulation. Steen held that this strip was made by palatine
and transverse bones, but the right lower jaw shows a coronoid process in position
with respect to the jugal and maxilla, which determines the anterior end of the
subtemporal fossa, and thus of the hinder end of the transverse bone ; it lies much
too far forward to make Dr. Steen's interpretation plausible. Furthermore, the
ridges of the ornament on the presumed " ectopterygoid " continue the direction
of those on the pterygoid, as they would be unlikely to do across a suture.

THE BRACHYOPID LABYRINTHODONTS 369

It is thus evident that Dr. Steen (and Dr. Romer) were unjustified in regarding this
skull as broad and low, and it remains to determine its shape.

As Steen recognized, the left premaxilla is in position and the series of teeth it
supports clearly shown. The parasphenoid — a little laterally bent — is in articulation
with the anterior part of the palate, and the dorsal surface of the skull — though the
sutures connecting its bones opened a little when it was spread out flat — is essentially
intact. It is evident that the part of the pterygoid which on the spectator's right
lies mesial of the basicranial articulation is the parotic plate, which in life was
essentially vertical.

Thus reconstruction of the skull shape is a matter of fitting measurements so that
dorsal, ventral and lateral views are self consistent and provide projected transverse
sections which give proper measurements. I began with the right maxilla, jugal and
quadratojugal, which show that the lower border of the cheek turns downward from
the direction of the straight maxillary border. The otic notch of the left side and
upper part of the squamosal in front of it are shown ; they allow the depth of the
cheek at its hinder border to be measured, and the size and shape of the right
quadratojugal and jugal fix the outline of the squamosal, though there is a possible
foreshortening by obliquity in direct side view. The width of the table, between the
otic notches, is shown for the left side, the interorbital width and the sizes and shapes
of the postfrontal, postorbital, intertemporal and supratemporal are known, and in
no Labyrinthodont is the table more than gently curved. Thus trial sections give
possible proportions. The skull shape so determined, with its very characteristic
down- turning of the lower border of the cheek behind the orbit, was then fitted to the
lateral aspect of the lower jaw as shown directly on the animal's right, the jaw being
shortened because it is a curved structure which has been laterally flattened into a
straight one.

The resulting fit is surprisingly good. Finally the palate has to be fitted in. The
distance measured from the most lateral point of the basitemporal facet of the ptery-
goid to its lateral border is known. The distance from the most dorsal point of the
parotic plate to the border of the pterygoid is known. The lowest possible place
for the outer border of the pterygoid with respect to the lower jaw is known. Thus
trial transverse sections made by measurements of individual bones along correctly
determined directions allow small modifications to be made until everything fits.
Finally the parasphenoid has to be put into place. The reconstruction of the skull
carried to the present point has fixed the position of the facet for articulation of the
pterygoid with the parasphenoid. The parasphenoid is still in position with respect
to the premaxilla. Direct measurements made on the skull photograph show that the
distance from the front point of the interpremaxillary suture to the articular facet
on the pterygoid agrees exactly with that for the pterygoid on the parasphenoid,
when the measurement is made on a trial longitudinal section of the skull. Thus
the slope of the bone in side view is established, the total length from the anterior
end of the skull to the hindmost point of the parasphenoid is known, so that the
position of the occiput is fixed with considerable certainty. The results are shown in
Text-fig. 24. It may be added that there are no traces of any cartilage bones in the
brain-case, so that the animal was probably very young.

37° THE BRACHYOPID LABYRINTHODONTS

PROBABLE RELATIONSHIPS OF PELION

As so restored Pelion has notable resemblances to Eobrachyops. The two
agree in having anteriorly placed orbits and a shallow face, behind which lies a long
post -orbital skull with a wide table, cheeks sloping steeply downward, a lower border
which passes downward from the hinder end of the short maxilla (at the level of the
hindmost point of the orbit) to a low-placed quadrate, and an occiput which slopes
backward below the hinder border of the table. The shape of the intertemporal is very
much the same in the two animals, which implies that the mutual relations of all the
bones in the postorbital region are very similar. In each the interpterygoid vacuity
is very large, and the palate deeply vaulted as a result of a great widening of the
pterygoid and the down-turning of its lateral strip mesial to the masticatory muscles.
Pelion and Eobrachyops have an identical mode of articulation between the hinder
surface of a processus internus of the pterygoid and a vertical facet on the laterally
widened anterior border of the parasphenoid. In each animal the lower jaw has a
very unusual straight lower border, and an upstanding coronoid process, not including
any contribution from the surangular, conditions otherwise unknown in Labyrin-
thodontia except in Dvinosaurus and the Brachyopids. But Pelion retains an otic
notch, a structure no doubt found in all primitive Labyrinthodonts, but lost in
Eobrachyops, Dvinosaurus and all other Brachyopids. Thus it is not improbable that
Pelion is closely related to the ancestry of Eobrachyops, and hence to that of Dvino-
saurus and the Brachyopids.

EOBRACHYOPS COMPARED WITH ERPETOSAURUS

My attention was called to Pelion in this connection by the existence in it of the
processus internus of the pterygoid, and, as Erpetosaurus has a similar structure,
that animal should also be reconsidered.

DESCRIPTION OF ERPETOSAURUS MATERIAL

Dr. Steen's material includes a skull (E. radiatus) in counterpart, so that an
imperfect dorsal surface (R.2672) and a wonderfully preserved hinder part of the
palate (R. 267oa) are available, together with an interclavicle, vertebrae and scutes
(R.2672). A second skull (E. laevis B.M.N.H., R.2662) shows a well-preserved
ventral surface, of which only the right hinder corner is missing, and another slab
(R.266o) contains an interclavicle, scutes and vertebral elements. I have rubber
casts of this material for comparison with Dr. Steen's descriptions, and can confirm
them, adding something because my casts are so much better than her squeezes.

No. R.2&7oa retains the impression of the hinder part of the palate of a skull whose
dorsal surface is shown by R.2&72. This skull has the lower jaw of each side
articulated with the quadrate, so that only small parts of this bone are visible. The
pterygoids, though broken into parts by cracks, are perfectly preserved in detail and
can be restored without difficulty. The ventral surface of the parasphenoid, and
left side of the brain-case — much dorsoventrally flattened but otherwise well

THE BRACHYOPID L AB YRINTHODONTS 371

preserved — are visible, but a short misplaced rib running antero-posteriorly below a
lappet belonging to the parasphenoid introduces some difficulties. The basioccipital
is perfectly preserved, uncrushed, in articulation with the parasphenoid. The right
exoccipital has swung round so that the whole of its posterior surface is shown,
whilst the left retains its natural position, but is dorso-ventrally compressed. The
parasphenoid is very well shown, and the left stapes is in its natural place.

Thus from enlarged photographs, on which the outlines of all important structures
are inked in, it is possible by trial and error to restore the whole occipital region,
measurements made directly from the casts with proportional compasses set at the
magnification of the photographs giving constant control and guidance. The results
are set out in Text-figs. 25 and 26, which in most matters explain themselves, but are
here supplemented by notes on some points.

BRAIN-CASE

Occipital condyle. The large basioccipital has a widened, trumpet-shaped hinder
end in front of which the bone narrows until it is surrounded by a ridge on the
hinder part of the parasphenoid, which overlaps it at a very obvious denticulated
suture. The " condylar surface " is a very deep conical pit. Each exoccipital bears
a large concave articular surface lying largely lateral to the foramen magnum,
joining the surface by which the exoccipital is attached to the basioccipital.

Occipital surface. The exoccipital is a very large thin sheet of bone of elaborate
shape. It is evidently mainly perichondral bone sheathing the paroccipital so
completely that it stretches far out beyond the vagal foramen, and wraps round the
lower surface of the otic capsule, mesial of the fenestra ovalis, so that its anterior
border has a long contact with the hinder border of the parasphenoid.

Supraoccipital. About one-half of the posterior surface of a supraoccipital showing
part of the border of the foramen magnum is seen in articulation with the right
exoccipital.

The parasphenoid is seen from below, a little hidden by a rib which is overlapped by
a free ventral lappet of parasphenoid, and partly concealed on the other (right) side
by a displaced ventral scute and fragments of pterygoid. The bone is cracked
transversely, the hinder part — when viewed from below — lying some half millimetre
dorsal to its original continuation. This is interpreted in Dr. Steen's restoration
(text-fig, i) as a pocket, but is shown by the new cast to be no more than a crack.
Behind this fracture the parasphenoid bears a wide median ridge whose ventral
surface becomes rounded posteriorly where it sheathes the basioccipital. The sides
of this ridge are almost vertical, but pass into the wide lateral expansions of the hinder
part of the bone. These pass backward until they meet the anterior borders of the
exoccipitals. Lateral to this contact the border passes outward and forward,
overlapping the prootic nearly to the border of the fenestra ovalis.

The paroccipital is represented only by a completely crushed but bony outer end,
lateral to the exoccipital. The left stapes is present in position. It is a powerful
bone with a wide flat foot, narrowing rapidly until it plunges under the quadrate

372

THE BRACHYOPID LABYRINTHODONTS

Tab.

TEXT-FIG. 25. — Erpetosaurus radiatus. Reconstruction of a skull (B.M.N.H.,
R.267oa). x a little more than 2. Dorsal aspect. (Text-fig. 26 is of the same
individual.) Reference letters as before.

THE BRACHYOPID LABYRINTHODONTS

373

Par.Oc.

TEXT-FIG. 26. — Erpetosaurus radiatus (B.M.N.H., R.26y2 and R.267oa). x a little more
than 2. Reconstructions A, of the hinder part of the palate, and B, occiput. Reference
letters as before.

374

THE BRACHYOPID L AB YRINTHODONTS

ramus of the pterygoid and is lost to sight. A very small perforation, presumably
for a stapedial artery, enters its ventral surface far laterally.

Tabular and dermosupraoccipital. These bones lie largely on the dorsal surface of
the table, but posteriorly they are deeply depressed where a pair of muscles spread
forward from the neck to overlap small rounded areas. These areas are bounded
anteriorly by a step rising to the highly ornamented dorsal surface of the skull, and are
largely visible from behind.

P. MX.

Vo.

Pal.

MX.

Ec.Pt.

TEXT-FIG. 27. — Erpetosaurus laevis (B.M.N.H., R.2662). X2j approx. Reconstruc-
tion of the palate. The dotted circle in the processus cultriformis of the parasphenoid
is the position of the pineal foramen. Reference letters as before.

PALATE

The palate as a whole is shown in E. laevis (R.2662), specifically distinguished by
Dr. Steen from R.267oa described above. The distinction is valid, the parasphenoid
of R.2662 lacking the free lappets of the other animal and bearing two grooves

THE BRACHYOPID LAB YRINTHODONTS 375

leading to foramina for the carotids, absent in R.267oa. They are, however, similar in
all important known characters. R.2662, though well-preserved and nearly complete,
is very difficult to interpret because the dentition is most unusual. The maxilla has
slid downward and outward over the palatine, and the great premaxillary teeth have
crushed forward and outward. Except anteriorly my figure agrees rather better
with Dr. Romer's than with Dr. Steen's, whilst the difficult premaxilla is unlike
either previous drawing.

VERTEBRAL ELEMENTS

The specimen R.2672 of Erpetosaurus has associated with the skull an interclavicle
and some other bones, together with some vertebral elements and very many scutes.
Dr. Steen has described and figured most of these, but the new cast makes it possible
to describe some vertebrae. A single intercentrum is seen end-on as a crescent-
shaped bone, exactly like that of Trimerorhachis. It is a short segment of a
cylinder of bone surrounding a notochord of large diameter and very little constricted.
Its visible semicircular border is sharp-edged and a little out-turned. It bears
no visible rib attachment. Closely associated are three pleurocentra. They are
exposed from their outer surfaces and, except for a transverse fracture of one, show
very little evidence of curvature to surround a notochord. They taper a little to
each end, and have part of the upper end of one margin everted so as to leave an
articular facet directed outward and either forward or backward. The two elements
of a single unfused neural arch agree, except that the spine of one stands across the
bedding and is crushed down and incompletely shown. The neural arch element has
a high spine widening as it passes upward ; its hinder border comes down on to the
postzygapophysis, whilst the anterior border is at the root of the anterior zygapo-
physis. There is no transverse process, or sign of rib articulation. Text-fig. 28, A,
is a restoration from these elements.

Another slab from Linton (B.M.N.H., R.266o) shows very many ventral scutes,
and part of the inner surface of an interclavicle, which resemble those of Erpetosaurus.
It is not capable of definite determination, but in many ways agrees with the original
(R.2664) of Steen (pi. 4, fig. 2) showing twenty-five or more of the " triangular plates "
which she figured. In this specimen the following matters seem certain (a) that all
the elements are paired, none fused into cresent-shaped intercentra, as is the single
example in R.2672 ; (b) all these bones have an internal surface toward the notochord
which appears granular, and a smooth outer surface ; (c) two types are present,
intercentra and pleurocentra. An intercentrum widens toward its lower border,
which turns in under the notochord but does not meet its fellow, and has a blunt
dorsal end. The hinder border is cut into by a rounded depression for surviving
cartilage continuous with the neural arch, the rib head being connected by a liga-
ment to a small pimple in the middle of the side of the bone. The other element,
a pleurocentrum, is somewhat deeper than the intercentrum, and narrows both
above and below. It bears an articular facet cutting into the summit of the
anterior border of its smooth outer surface. The neural arch is very poorly shown,
though there is an example in articulation with an intercentrum and pleurocentrum
(Text-fig. 28, B) ; in no case is there any sign of a transverse process. I can only

376 . THE BRACHYOPID LAB YRINTHODONTS

assume that here and in Erpetosaurus the rib articulated with the neural arch and
pleurocentrum where the facet existed, and that in the present specimen its other
head was tied to the " pimple " without any contact. There is evidence that the
rib forks, one of the " heads " having a pointed, non-articular tip. The differences
from the Erpetosaurus vertebrae already described may well be due to differences
along the length of the column.

Thus the structure of Erpetosaurus is relatively well known, and its systematic
position should be easily determinable. The vertebral column, as Dr. Steen has
shown, agrees with that of the contemporary Stegops, and is of a type found in the
Middle Rothliegende Acanthostoma, where as she states (1937 : 499), " both pleura-
and intercentra being ossified as right and left hemicylinders, unfused in the mid-
dorsal or mid- ventral line." Romer (1947 : 169) doubts this interpretation, suggest-
ing that the intercentra shown in Steen (1937, text-figs. 4 and 5) are twisted round

B

TEXT-FIG. 28. — A, Erpetosaurus radiatus (B.M.N.H., R.26y2). Reconstruction from
scattered elements of three vertebrae from the right side. The intercentra are single ;
B, ? Erpetosaurus (B.M.N.H., R.2664). Three vertebrae and a rib from the right.
The shaded set of elements is drawn as it actually occurs. The intercentra are paired.
All about 2 diameters.

and seen from below. I have examined casts of the specimen and its counterpart,
which are not good enough to permit any definite conclusion, but Dr. Steen, with the
original slabs before her, is unlikely to have been wrong in such a point.

Thus these vertebrae differ from ordinary rhachitomous vertebrae (a) in the absence
of a transverse process well separated from the parachordal part of the neural arch,
(b) in the usual existence of paired intercentra, which sheath the notochord for a
large part of its circumference, (c) in the very high pleurocentra which extend over a
very large part of the notochord. They are what Romer (1947 : 64, fig. n) calls
protorhachitomous, which by further ossification could well become embolomerous.
Stegops is supposed by Romer to be related to Acanthostoma, which is evidently close
to Zatrachys. If these views be justified then we have for the first time direct

THE BRACHYOPID LAB YRINTHODONTS 377

evidence of the evolution of typical or nearly typical rhachitomous vertebrae from
such a protorhachitomous type.

SIMILARITIES BETWEEN THE Two FORMS

Thus, except for the processus internus of the pterygoid and its mode of articulation
with the parasphenoid, there is in Erpetosaurus no resemblance to Eobrachyops such as
that which Pelion shows ; but the vertebrae of Eobrachyops are " protorhachitomous",
comparable with those of Erpetosaurus and Stegops.

PLATYSTEGOS LORICATUM Dawson

DESCRIPTION OF MATERIAL

Another, earlier, Labyrinthodont, with a processus internus as a mode of
articulation of the pterygoid with the parasphenoid, is an incomplete skull, apparently
referable to Platystegos loricatum Dawson, from the Coal Measures of South Joggins,
Nova Scotia, described by Dr. Steen (1934 : 480, 481). The South Joggins section is
thick — 11,000 ft. according to Dawson — and the amphibia come from the interior
of hollow lepidodendroid trunks which stand on coal seams. Most are from
Division IV, Section XV of the section ; this is about half-way up. The evidence of
age comes mainly from plants, but according to Bell (1944) they are Westphalian B.
Bell noted that the whole series does not extend high enough to reach the first
occurrences of Anthraconauta phillipsi, but contains molluscs compared to Anthra-
comya cf. modiolaris. Thus the series seems to correspond to the Middle Coal
Measures of Staffordshire, and the horizon of Platystegos, though above that of
Eugyrinus wildi, is well below that of Linton and Nyrany.

This skull (D.M.S.W., 6.95) shows the lower surface of a skull table with the
upper part of the right orbit and the face in front of it. The block containing this
complex of bones fits on to two others, one of which retains a mould of the lower
surface of the table and the hinder parts of the two pterygoids, the left essentially in
natural position though forced a little (perhaps 3 mm.) upward, the right misplaced.
The third block continues the latter and shows the outer surface of the left side of the
skull from premaxilla to quadrate, the quadrat ojugal being misplaced and lying in
the orbit. The right pterygoid extends forward until it overlaps the vomer and
palatal bones, whose upper surfaces are shown. The jugal of this side is detached
and visible from nearly all aspects. The brain-case is completely missing, but one
ramus of the lower jaw is largely seen.

STRUCTURE OF SKULL

The general structure is clearly shown in the drawings in Text-figs. 29 and 30
which, though restorations, are carefully measured projections based on adequate,
undistorted remains. The skull is short and broad, a little less than half its maximum
breadth in height. The orbits are large and in the middle of the length, and the
otic incision is overhung by the tabular, which forms a short process behind it. The

378

THE BRACHYOPID L AB YRI NTHODONTS

hinder border of the dermosupraoccipitals and tabulars is turned downward, and
there is some evidence that the occiput extended considerably behind the table and
the quadrate condyles. There is a well-defined scar on the inner surface of that
ridge of the tabular which forms the upper border of the tympanic notch which can
only have articulated with the end of the paroccipital process. It is thus evident
that no process passed inward from the tabular in contact with the upper surface of

I. No. 0P.

Lac.

I. Tern.

S.Tem.

Tab.

S.Tam.

TEXT-FIG. 29. — Skull of Platystegos loricatum Dawson. (D.M.S.W., 6.95) x ij.
Restored drawings of the skull. The sutures on the skull roof are taken from the lower
surface. A, Dorsal aspect ; B, right side. Art.Proc.Sq., articular process of the
squamosal for attachment to the supratemporal ; D.S.Oc., dermosupraoccipital ;
I.Na.Op., internarial opening ; I .Tern., intertemporal ; Lac., lachrymal ; Qu.,
quadrate ; S.Tem., supratemporal ; Sq., squamosal ; Tab., tabular.

THE BRACHYOPID LAB YRINTHODONTS

379

the paroccipital and below the post-temporal fossa. This condition exists in the
Loxommids, but has been lost in all other Labyrinthodonts. There is an inter-
temporal bone separating the supratemporal from the postfrontal, and the lachrymal
— which shows the course of the duct with its two canaliculi from the orbit — forms a
large part of the orbital margin and the hinder border of the nostril.

The squamosal is remarkable because it is not attached suturally to the outer
margin of the supratemporal. It abuts against it, the bone being 1-5 mm. — or a
little more — in thickness in a skull some 5 cm. long. To strengthen this attachment

Vo

Pal.

D.S.Oc.

Tab.

TEXT-FIG. 30. — Skull of Platystegos as in Text-fig. 29. x i£. Palate. D.S.Oc.,
dermosupraoccipital ; Ec.Pt., ectopterygoid ; Ju., jugal ; Pal., palatine ; Proc.Int.Pt.,
processus internus of the pterygoid ; Pt., pterygoid ; Qu.R.Pt., quadrate ramus of ptery-
goid ; Tab., tabular ; Vo., vomer ; * pterygoid process described in text, p. 381.

a special process arises from the upper end of the admesial border of the subotic
flange of the squamosal ; this process passes directly inward with its essentially flat
but dimpled upper surface opposed to a special sharply marked depression on the
lower surface of the supratemporal. This arrangement, which is seen on both sides
of the specimen, implies a powerful, ligamentous attachment.

The left quadrate is well shown from behind. Its lateral border is notched imme-
diately above the shallow but wide outer condyle by the quadratojugal foramen, and
a narrow parallel-sided strip of its posterior surface extends upward, visible between
the inner border of the sub-tympanic flange of the squamosal and the hinder border
of the parotic flange of the pterygoid. Some distance above the bony quadrate the

GBOL. II, 8. 32

380

THE BRACHYOPID LAB YRINTHODONTS

inner edge of the squamosal steps inward to meet the pterygoid, presumably above a
cartilaginous upward extension of the quadrate. There is a considerable pterygoid
flange of the quadrate.

The pterygoid is a remarkable bone, not to be matched in any other Labyrin-
thodont known to me. The attachment to the basis cranii is by a processus internus,
whose lower surface is rounded from back to front. This is abruptly truncated by a
face which evidently articulated with a flat transverse facet on the parasphenoid.
The anterior margin of the process begins at a point mesially and bows forward to its

T?b- D.S.Oc.

S.Tc

Arh Proe.

TEXT-FIG. 31. — Skull of Platystegos as in Text-fig. 29. x ij. A, Posterior view of the
skull, lacking brain-case ; B, the posterior part of the skull, without brain-case,
cut across about at the middle of the supratemporal and viewed from in front.
Cut surfaces are left white. Avt.Proc.Sq., process of the squamosal which is applied to
the supratemporal ; D.S.Oc., dermosupraoccipital ; Par., parietal ; Proc.Int.Pt.,
processus internus of the pterygoid ; Pt., pterygoid ; Pt.No., " notch " in the
pterygoid ; Qu., quadrate ; Qu.Cart., cartilaginous continuation of the quadrate ;
Qu.J., quadratojugal ; S.Tem., supratemporal ; Sq., squamosal ; Tab., tabular ;
* pterygoid process described in text, p. 381.

root. It there swings forward to continue as the border of a rather large inter-
pterygoid vacuity. This border can be seen on the right side until it passes below a
misplaced lower jaw and cannot be followed. But it is evident that the pterygoid is
still wide when it reaches the vomer, and that it may well have reached the para-

THE BRACHYOPID LAB YRINTHODONTS 381

sphenoid. The lower surface of the processus interims continues laterally without
break into that of the body of the pterygoid, which lies mesial of the vacuity for the
masticatory muscles, and extends down within them to a curved ventral border.

The processus internus lies so far back in the skull that the quadrate ramus is not
distinguishable from the body of the bone. The ramus sheathes the pterygoid
process of the quadrate, rolling over its ventral border so as to form a rounded
transition from the horizontal body to the concave upper part of the quadrate ramus.
The quadrate ramus thus sheathes the ventral border and posterior surface of the
quadrate, extending forward and inward beyond that bone to rise to the skull roof,
behind the articular process of the squamosal. This transversely placed part of the
pterygoid has a free admesial border, whose lower end (marked * in Text-figs. 30, 31)
— lying above and a little behind the processus internus — projects backward to a
point not meeting any other bone. The posterior surface of the pterygoid, between
this point, the skull roof and the quadrate, is a hollow cone, incomplete because it
lacks a triangular strip of its ventral part between the quadrate and point *. That
flange of the pterygoid which sheathes the temporal muscle ends anteriorly by its
lower border rising abruptly to pass into the lateral border of the palatal part of the
bone. It comes, if not into direct contact with, at least into the very near neigh-
bourhood of, that internal process of the jugal which, as in the majority of Labyrin-
thodonts, lies dorsal to the ectopterygoid, and often extends behind it to form the
anterior border of the subtemporal fossa. The whole palatal surface of the pterygoid
seems to have been covered by a sparse granulation of small teeth, little more than
hemispherical projections.

No definite part of the ectopterygoid is visible, but the anterior end of the right
palatine is well shown from above and from the outer side. The admesial part of the
bone is a thin sheet with a feather edge. This passes laterally into an upturned ridge
whose lateral face is deep, rounded from dorsal to ventral borders, and longitudinally
grooved, evidently in continuous contact with the maxilla. This border ends by
sloping down as the upper surface of a deep notch, the hinder border of the palatal
nostril. Immediately below this process the bone carried a tusk, circular in section,
with a sharply-grooved base. The large vomer is overlapped by the anterior end of
the palatine, mesial and anterior to which it has a wavy thin posterior border, presum-
ably articulating with the pterygoid. It is evident that some part of the bone has a
granulation of small blunt teeth, widely spaced, and similar to those of the pterygoid.
There is evidence of a vomerine tusk placed perhaps mesial of and near to the anterior
end of the palatal nostril.

The left premaxilla is present in place, attached to the admesial end of the left
maxilla. The anterior part of the internasal opening is shown. The suture with
the nasal and the bone's contribution to the border of the nostril are uncertain.
Some of the premaxillary and maxillary teeth are shown, and justify the figures.
The general shape is certain, as the left side of the skull from front to back is visible,
giving no evidence of distortion except for an inturning of the maxillary lower
border, and the other drawings are fitted into the space so determined.

The inner surface of the hinder end of the right ramus of the lower jaw
is shown from above, and an irregular transverse section of its anterior part

382 THE BRACHYOPID L AB YRINTHODONTS

is shown on a fractured surface. The face on the articular for the quadrate is
fully ossified, divided by a low ridge into inner and outer regions for the quadrate
condyle. It is clasped by the surangular laterally, and the prearticular — which
sheathes its inner surface and continues behind it to meet the surangular — forms with
that bone a ridge on the posterior surface, penetrated by a circular canal presumably
for the chorda tympani. The rounded bottom of a groove on the prearticular helps
to isolate the ridge, which is unfortunately damaged by a chisel cut, but may have
been a first beginning of a retroarticular process. In front of the condyle the upper
border of the prearticular descends very rapidly so as to leave the cavity of the jaw
widely exposed from within. Anteriorly, below the orbit, the jaw becomes shallow,
nearly as wide as it is high, with a marginal row of teeth bounding a flat upper surface.
Thus it is possible that the Labyrinthodont jaws from South Joggins, recognized by
Dr. Steen (1934 : 487) as those which first exhibit the later Labyrinthodont retroar-
ticular process, belong to Platystegos.

COMPARISON WITH DENDRERPETON AND EUGYRINUS

Platystegos is found with Dendrerpeton and a series of very similar forms described
by Dawson and Dr. Steen. The proportions, safely determined in each case, show
that they may reasonably be generically distinguished. But the two animals are very
similar ; the pattern of the external skull bones is essentially the same, the general
nature of the palate with its large interpterygoid vacuities apparently bounded by
the pterygoids and parasphenoids alone is not too dissimilar, and a processus internus
of the pterygoid articulates with the parasphenoid in each. In each the tabular
has a descending process from its under surface behind the tympanic space, and the
general outline of the ventral borders of the descending flanges of the tabulars and
dermosupraoccipitals is similar. In each form, if we may judge Dendrerpeton by
Steen's photograph (pi. i, fig. 2) the subotic flange of the squamosal passes smoothly
over a rounded surface to the outer side of the skull, there being no well-defined
crest as a border to which the tympanic membrane could have been attached. The
pterygoid of Dendrerpeton is not known to have the extraordinary conical form of the
quadrate ramus which occurs in Platystegos, but it extends up to the skull roof (Steen,
text-fig. 2g) in the same way, and there is evidence in B.M.N.H., R-4553 and R.4554
that the special process of the squamosal which supplements the straight abutment
of that bone on the table in Platystegos actually occurs in much the same form.

The lack of an interdigitated suture between the squamosal and the skull table in
Dendrerpetonts, whilst all other dermal skull bones are firmly attached in that
manner, seems to show that the contact itself is of recent introduction into their
structure, the development of the special articular process of the squamosal showing
that a firm connection is desirable. Similarly in Anthracosaurs the squamosal
alone of dermal skull bones fails to have a sutural attachment to the table, and is
held in place by the intervention of the palatoquadrate cartilage. The only explana-
tion of the long delay in the establishment of this connection so implied is the survival
in their Osteolepid ancestors to a relatively recent period of the notch between the
membrane bones of the cheek and table which existed in them.

THE BRACHYOPID L AB YRINTHODONTS 383

The group of Dendrerpetonts is additionally characterized by the occurrence of
bony scales, oval or shield-shaped, with a delicate concentric striation, together with
faint radial striae. In many ways Platystegos recalls Eugyrinus (Watson, 1940 : 215)
which is somewhat older and much smaller. The general pattern of the dermal skull
bones is common to the two, though Eugyrinus possesses lateral line grooves,
apparently absent in all Dendrerpetonts. Both have large occipital flanges from
the tabulars and dermosupraoccipitals, and the large parotic flange of the pterygoid
reaches the skull roof behind a special articular process of the squamosal in the same
way. The mode of articulation of the pterygoid with the basis cranii is different,
Eugyrinus not possessing a typical " internal process ", and rather resembling
Eryops. Eugyrinus has a " pterygoid flange " against the lower jaw absent in
Platystegos. The dermal scales of Eugyrinus have a general resemblance to those of
Dendrerpetonts. But in their possession of large interpterygoid vacuities the two
groups agree, differing very greatly from the contemporary Loxommids.

COMPARISON WITH BRACHYOPIDS

Eobrachyops is, for its time, of very advanced and specialized structure. It retains
an intertemporal bone and has thus a skull pattern of normal early Labyrinthodont
type. But its face is extremely flattened, and its temporal region deepened im-
mensely. The mechanical effect is to enable the animal to open its mouth to an
extraordinary extent, by the action of a museums depressor mandibuli attached to a
retroarticular process much longer than in any contemporary. The long temporalis
muscle would secure a powerful snapping bite. The wide and deep arch along the
palate, between the pterygoid flanges which pass down mesially to the temporal
muscle, would enable a large prey to be swallowed and could house a large tongue.
The sloping occiput is associated with a very posteriorly-placed condyle, and perhaps
with the possibility of raising the head nearly vertical to the vertebral column, and
the considerable area of membrane bone on the occiput provides for the attachment
of powerful neck muscles.

The stapes of Brachyopids, known in Batrachosuchus and Dvinosaurus, is presum-
ably a modification of the normal Labyrinthodont type, subsequent to the suppression
of the tympanic membrane which must have existed in their ancestors if they were to
possess a stapes at all. This loss of an otic notch is associated with a movement back
of the point of contact of the squamosal with the skull roof, and involves a correspond-
ing backgrowth of the parotic part of the pterygoid. In association with this
change in shape a horizontal flange, essentially a development of the base of the
processus internus, comes to arise abruptly from the inner surface of the pterygoid ;
separating the parotic plate from that which sheathes the masticatory muscles
almost down to their attachment to the mandible.

The quadrate is unique. Its flat inner surface is completely covered by the
pterygoid, but the bone itself is exposed as a narrow vertical rib between the ptery-
goid and the admesial border of the squamosal. Laterally to the quadrate rib the
squamosal forms a vertical hollow cylindrical face before passing over a rounded
surface to the cheek.

384 THE BRACHYOPID LAB YRINTHODONTS

Inspection of the figures of Platystegos will show that in many ways it makes a
starting point from which the Brachyopids could have arisen. A deep ridge of
quadrate is visible from behind between the pterygoid and squamosal, which turns
a very little backward towards it. The posterior face is widely and shallowly
grooved from top to bottom and it passes over a rounded surface to the outer side,
there being no sharply defined ridge for the attachment of a tympanic membrane.
The conditions differ from those in Brachyopids in the retention of a definite pterygoid
ramus of the quadrate and of the pterygoid sheathing it. But these are common
early Labyrinthodont features, which must certainly have existed in some ancestral
Brachyopid.

The palate of Platystegos has, in the deep flange of the pterygoid which sheathes the
masticatory muscles, a structure which alone could have provided the starting point
of the Brachyopid condition. The mode of articulation of the pterygoid with the
parasphenoid is the same in Platystegos and Eobrachyops, and the large interpterygoid
vacuities provide conditions from which the relatively enormous vacuity of Eobra-
chyops could have arisen long before it is parallelled in most other Labyrinthodonts.
It is, of course, most improbable that Platystegos is a real ancestor of the Brachyopids :
all that can be claimed for it is that in some ways it possesses structural qualities
which explain how those which characterize that group may have arisen. On the
other hand Platystegos differs as completely as any Labyrinthodont can from
Erpetosaurus, which seems to have been a rapidly advanced form with no known
descendants.

Platystegos is also important because with its associated Dendrerpetonts and
Eugyrinus it is the earliest known normal Labyrinthodont, other than the Loxommids,
which retains obviously more primitive features in many structures in the skull,
together with certain special peculiarities, and an advance in the mode of attachment
of the squamosal to the table.

/

SURVEY OF BRACHYOPID LINE

We have thus by consideration of the structure of skulls (and lower jaws when
they are known) reached the conclusion that Eobrachyops townendae, Clear Fork ;
Eobrachyops casei, Clear Fork ; Bothriceps australis, ? M. Permian ; Dvinosaurus,
U. Permian ; Trucheosaurus, U. Permian ; Brachyops, Permo-Triassic boundary ;
Batrachosuchus browni, L. Trias; Batrachosuchus watsoni, L. Trias; " Platyceps"
wilkinsoni, L. Trias ; Plagiosternum granulosum, M. Trias ; Plagiosuchus, M.
Trias ; Gerrothorax, U. Trias and Rhaetic ; and Plagiosaurus depressus, ? Rhaetic ;
are members of a group of Labyrinthodonts relatively closely allied to one another,
though not members of a single lineage. These come from Texas, Australia, Russia,
India, South Africa, Germany and Sweden, and members of the group are known
also from Argentina, Siberia and Spitsbergen. But ten species are known by a
single specimen only, two from about three specimens each, and only Dvinosaurus is
represented by a satisfactory suite of some seventeen. Such a mode of occurrence
suggests that the evolutionary history of the group was complex, and the occasional
recognizable existence of neoteny emphasises the probability. Furthermore, nothing

THE BRACHYOPID L AB YRINTHODONTS 385

in the known structure of a single young skull referred to Pelion from the Coal
Measures of Ohio prevents its acceptance as an early member of the group, but it
differs in some ways which will require discussion.

Thus it is evidently impossible to hope to find a true evolutionary series amongst
these animals ; the whole group would probably have included a large number of
species representing not only members of main stocks, but many blind-ended side-
branches like Dvinosaurus, retaining, when sexually mature, larval qualities which
may be (or may, like a frog's tadpole, not be) proper to a time earlier than that in
which they lived.

Thus all we can do is to study the group as a whole, placing its members in order
of time and recognizing such changes in structure as persist over all or some part
of the long life of the stock. I did this (so far as the material allowed) in 1919
(p. 56) showing that the direction of change in structure was the same as in the
Capitosaur-shaped skulls from Eryops to Cyclotosaums.

The greatly extended knowledge contained in the present paper confirms that
conclusion and extends it :

(1) The large mainly basioccipital condyle of Eobrachyops passes to the wide,
tripartite condyle of Dvinosaurus, and that passes to the separated exoccipital
condyles and absence of a basioccipital in Batrachosuchus, etc.

(2) The ossified supraoccipital of Eobrachyops and Dvinosaurus vanishes in the later
forms.

(3) The widespread exoccipital of Eobrachyops, although reaching the tabular,
does not extend forward to the pterygoid as it does to an increasing extent in
Bothriceps, Batrachosuchus and the " Plagiosaurs ". The otic capsule contains no
visible ossification in any form.

(4) The pterygoid, movably articulated with the parasphenoid in Eobrachyops
and Dvinosaurus, gains a sutural attachment to the lateral border of the para-
sphenoid in Bothriceps, which steadily increases in extent by spreading backward
to the exoccipital.

(5) Hypoglossal foramina perforate the exoccipital of Eobrachyops, lie on the lateral
surface of that bone in Bothriceps, emerge through the hinder surface in Batracho-
suchus, and have disappeared in Gerrothorax.

(6) The occipital aspect of the skull of Eobrachyops shows a deep occiput and very
ventrally placed quadrate condyles. Bothriceps, Batrachosuchus and Gerrothorax
show a progressive flattening, both of the lateral suspensory parts of the skull and of
the occiput.

(7) The quadrate, which in Eobrachyops is on the level of the occipital condyle when
the skull is viewed laterally, moves forward in late forms like Batrachosuchus and
Gerrothorax.

(8) The interpterygoid vacuities of Eobrachyops are of very great size, and there
is no real possibility of their increase in later members of the group.

(9) The retroarticular process of the lower jaw, present in Eobrachyops, considerable
in Bothriceps, becomes large in Batrachosuchus and immense in Plagiosuchus.

(10) The intertemporal, present as an independent bone in Eobrachyops, fuses with
the postorbital in Dvinosaurus and vanishes in Batrachosuchus and all later forms,

386

THE BRACHYOPID LAB YRINTHODONTS

TEXT-FIG. 32. — A series of Brachyopid skulls reduced to the same width. Various
magnifications. Dorsal aspects of A, Eobrachyops townendae, L. Permian ; B, Dvino-
saurus, U. Permian ; c, Bothriceps australis, ? M. Permian ; D, Brachyops, Permo-
Triassic boundary ; E, Batrachosuchus browni, L. Trias ; F, Gerrothorax pulcherrimus,
U. Trias.

THE BRACHYOPID LAB YRINTHODONTS

387

TEXT-FIG. 33. — A series of Brachyopid skulls reduced to the same width, various magni-
fications. Palatal views of A, Eobrachyops townendae ; B, Dvinosaurus ; c, Bothri-
ceps ; D, Batrachosuchus watsoni. E-F, Dorsal views of E, " Platyceps " wilkinsoni ;
F, Batrachosuchus watsoni.

388

THE BRACHYOPID LAB Y RINTHODONTS

the supratemporal thus coming into contact with the postfrontal, as in rhachitomous
and stereospondylous Labyrinthodonts in general.

(u) Eobrachyops has protorhachitomous vertebrae, in Dvinosaurus they are
rhachitomous, though the first vertebra lacks a pleurocentrum, and that of the next

B

TEXT-FIG. 34. — A series of Brachyopid skulls reduced to the same width, various magni-
fications. Left-hand column lateral aspects, right-hand column occipital views.
A, Eobrachyops townendae ; B, Bothriceps australis ; c, Brachyops ; D, Batrachosuchus
watsoni ; E, Gerrothorax pulcherrimus .

two fuses with the intercentrum and neural arch. In Plagiosaurs the neural arch
and intercentrum alone remain, the latter bone taking on the appearance of a centrum.

Thus in all important matters the course of evolutionary change in Brachyopids is
uniform in direction over the immensely long history of the group, and is the same
fundamentally as that found in the Capitosaur relatives.

This paper was begun during the war years, carried further in the late 1940*5 and

THE BRACHYOPID L AB YRINTHODONTS 389

finally written in University College in 1954. All the drawings, which are recon-
structions based on careful measurements made from adequate materials, were made
by Miss J. Townend after detailed discussions, extending over very long periods, of
the significance of all visible details, and the text owes much to her suggestions.

I am indebted to the Royal Society for enabling Miss Townend to work with me,
and to University College, London, and Professor Medawar for their hospitality to
Miss Townend and myself. The departmental staff deserve our thanks, and we owe
much to the splendid photographs made by Mr. W. H. Brackenbury on which our
restorations are often based.

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2 MA!) 1956

EXPLANATION OF PLATE 39

" Platyceps " wilkinsoni Stephens, type specimen, x 3. Narrabeen beds, Gosford, New South
Wales. Property of the N.S.W. Geological Survey, Sydney, Australia. (Compare outline
Text-fig, n.)

Hull. B.M. (X.H.) GeoL 2, 8.

P L A T E

#''

- 2 MAS 1956

INDEX TO VOLUME II

New taxonomic names and the page numbers of the principal references are printed in Clarendon type.

An asterisk (*) indicates a figure.

Abies .

Acanthostega .

Acanthostoma

Acheloma casei

A cinonyx pardinensis

Adelospondyli

Adiantites bassleri .

— bellidulus .

— peruianus

— whitei
Agathis
Alces

— dices

— latifrons .
Alcinae

126

320, 329
320, 376

• 355. 356
86

. 329

199, 200, 203, 209, 214
196

199, 202, 203, 214

199, 202, 203

121, 126, 129, 134

16, 19, 22, 61, 82, 83

14, 16, 25*, 83

. l6, 19, 22, 27, 46

7, 8, 10, 22, 26, 90

Anthraconauta phillipsi .
Anthracosauria,
Antlers, development of .
Aphlebia australis .
ARAMBOURG, C.

• 377

320, 329, 382

11-14

199, 200, 203
249

American Museum of Natural History 344, 356
Amphiprox . . . . . .13

Anemia ...... 293

— adiantifolia . . . 294, 303, 306

— affinis ...... 306

— anthriscifolia ..... 306

— aurita ...... 306

— colwellensis 291-293, 298*. 299, 304, 305-
307 ; PI. 35, figs. 39, 40, 42-53 ; PI. 36, figs.

59-64

— dregeana ...... 294

— ferruginea ..... 306

— fremonti ...... 294

— — forma fertilis . . 294, 303, 306

— fulva ...... 306

— gardneri 302, 303, 306; PI. 32, figs. 11-13

— imbricata ...... 306

— lanipes . . . 294, 302, 303, 306

— perrieriana . . . 302, 303, 306

— phyllitides . . 300, 302, 303, 306

— poolensis 291, 292, 294, 295, 296*, 297, 298*,
299, 302, 303, 306 ; Pis. 32-34 ; PI. 35, fig.
41 ; PI. 36, figs. 54-58

— schimperiana 294, 302, 303, 306 ; PI. 33,
ng. 23

— sp. 3 305, 307 ; PI. 36, fig- 65

— subcretacea . . 291, 292, 293, 294

— tomentosa ...... 306

Anisopteris . . . . . .195

Anoglochis . . . . . .11

Anonaceae . . . . . .165

Anonaspermum . . . .152, 165

— aegypticum 151, 165, 166 ; PI. n, figs. 13-15

— commune ..... 165, 166

— pulchrum ...... 166

— rotundatum . . . . .166

— rugosum ...... 166

Anthracomya cf. modiolaris . . -377

Araucaria 99, 109, 112, 116-118, 121, 122, 124,
126, 128-131, 133, 134

— angustifolia . . . 99, 104, 118, 131

— araucana .... 99, 104, 118, 131

— bidwilli .... 116-118, 131

— brasiliana .... 99, 131

— excelsa ...... 103

— grandifolia . . . . .104

— grandis . . . . . .132

— imbricata . . . . . 99, 109
- mirabilis 99, 107, 108*. 110, 113*, 114*,

115*, 118, 120, 131-134; Pis. 3-5
seedlings . . 107, 108*. 134 ; PI. 2

— pichileufensis . . . . .132

— windhauseni . . . . .no
Araucariaceae . . 106, 121, 122, 134
Araucarian galls . . . . .109
Araucarioxylon . . . . .106
Araucarites .... 102-105, 118

— mirabilis . . . . . .no

— patagonica . . . . .131

— phillipsi no

— sanctaecrucis 102, 105, 118; PI. i, figs, i,
4-6, 8-13

Archaeocalamites radiatus . . . 192

Archaeopteridium dawsoni . . .216
Archaeopteris cf. roemeriana . . .216
Archaeosigillaria . . . . .217

— vanuxemi . . . . . .217

Archegosaurus .... 344. 366

Archeria ...... 364

Arctodendron . . . . .217

Argentina (localities) 200, 201, 212, 213, 321,

323. 329, 384
ARNOULD, M. ..... 249

Arvicola ...... 87

— bactonensis ..... 86

— greeni ...... 86

— mosbachensis ..... 86
Asolanus ...... 194

— ? minimus . . 198, 202, 206, 208
A splenites prae-allosuroides . . 308, 309
Asplenium sp. . . . . . 307

— subcretaceum ..... 293
Asterocalamites . . . . 195, 197

— scrobiculatus . . . . .217
Athrotaxis . . . . 130, 132, 133

— ungeri . . . . . .132

394

INDEX

Australia

Auversian

Axis

AZZAROLI, A.

318, 321, 322, 329, 384

292, 311

II

. 1-96

Bagshot Beds . . . 291, 292, 309

Baiera pluripartita . . . .191

BAIRD, D. ...... 366

Barcombe Mills, skulls from . 229, 260, 268
Basel Museum . . . . 24, 45

Batrachosuchus 321, 323, 329, 331, 337, 341-
343. 366, 383, 385

— browni 321, 322, 331, 335*, 338, 338, 384.
386*

— ? sp 336, 337*. 338*

— watsoni 323, 331, 332*, 333*. 335, 336, 338,
384, 387*. 388*

BAYNES-COPE, A. D. . . 262, 283, 284
Benthosuchus ..... 336

Bessarabia . . . . . . 8, 9

Bison ....... 87

— priscus ...... 86

Blastocerus . . . . . 14,15

BLOOMFIELD, C. . . . 252, 255, 268

BOLTON, H. L. . . . . . 248

Bone implement .... 246, 253

Bones, composition of . . . 262
Bos 87

— primigenius ..... 86
Bothriceps . . 321, 329, 337, 385, 387*

— australis 321-828, 324*, 325*, 384, 386*,
388*

— major .... 322, 323, 327
Bothrodendron . . . 198, 205, 206

— australe ...... 201

— kiltorkense . . . . .196

— leslii ...... 205

— pacificum .... 198, 202, 204

— sp. . . . . 193, 194, 198, 202, 204
Boulder Clay ..... 5
Bournemouth Beds . 292, 293, 309, 310
Bovidae, distribution of . . . .87
BOWIE, S. H. U. . . . 262, 276-282
Brachyopidae 321, 322, 323, 328, 341, 342, 365,

366, 383, 384, 388
Brachyops . 322, 323, 329, 384, 386*, 388*

— laticeps . . . .321, 322, 330*
Brachyoxylon . . . . .106
Brachyphyllum . . . . .105

— mammillare . . . . .105
Brachyprosopus vireti .... 86
BRACKENBURY, W. H. . . . . 389
Branchiosauravus ..... 367
Branchiosaurs .... 320,344
Bunya (Araucaria) 99, 104, 109, 116, 118, 124,

131. 132, 134

Calamites

— peruvianus

— suckowi

— undulatus
Calamostachys sp.
Calathiops

— cf. plauensis
CALDER, M. G.
Callixylon

191, 193, 197, 201, 203

197, 199, 200, 201

191, 192, 195, 197, 201

195. *97. 201, 203

192

... . 216

. 217

97-138

216

Canada

Canine, Piltdown
Capitosaurus .
Capreolinae .
Capreolus

— australis .

— caprea

— capreolus .

— mut. prisca

— ruscinensis
Carboniferous
Cardiopteridium
Cardiopteris .
Carhaumayo flora .

318,329

. 239, 240, 256, 257, 271

326, 334. 330, 350. 363, 388

. 7, 90

30, 90

10

32

30, 32, 88, 90

30

10

189-224, 329, 343

201
. 216

209, 215

Carpolithus hassani 151, 183 ; PI. 16, figs. 64, 65

— sp. . . 151, 184 ; PI. 15, figs. 55-57

(Icacinicarya) 151, 184; PI. 16, fig. 66

Castor ....... 86

— fiber . . . 252, 253, 263, 269, 279
Cerro Cuadrado, Patagonia ... 99
Cervalces ...... 90

— scotti 19. 26

Cervavitus ...... 9

— demissus ...... 10

— tarakliensis ..... 8
Cervid indet. .... 77*. 78*
Cervidae, phylogeny of . . . 7
Cervinae . . . 7, 8, 10, 11, 47, 48, 90
Cervocerus ...... 9

— bessarabiae ..... 9

— novorossiae . . . . 8, 9, 9°
Cervus . . . n, 32, 34, 35, 47, 48, 90 ,91

— acoronatus .... 34, 35, 86

— albirostris . . . . .32

— ardeus ...... 45

— belgrandi ... • 4. 53. 54

— benindei ..... 34, 35, 86

— canadensis ..... 32

— capreolus ...... 32

— carnutorum . . . . 45, 83

— cashmirianus ..... 32

— cazioti ...... 67

— ctenoides ..... 37, 38, 40

— dama polignacus .... 83

— dama priscus ..... 52

— dawkinsi ..... 53> 61

— dicranios ...... 37

— elaphus 32-34, 35, 36*, 37*, 76, 87, 88, 90,
253. 263, 269, 279

angulatus .... 35, 88

atlanticus 33

barburus 33

bolivari 33

brauneri 33

corsicanus ..... 33

hippelaphus 33

hispanicus 33

maral ...... 33

— — mauerensis ..... 34

— — mosbachensis . . . -34

priscus . . . . 34. 35

scoticus 33

— etueriarum ..... 79

— falconeri ...... 37

— fitchii 61

— gunni ...... 61

— hanglu 32, 33

— /art/irons ...... 27

INDEX

395

Cervus macneilli

32

— maral fossilis

35

— martialis .

53

— megaceros

83

— obscurus .

73, 74*,' 75*. 76*. 78,

79.

89-91

— orticeros

37

— perolensis

ii

, 81

— philisi

ii, 81

. 86

— pliotarandoides .

47

— polignacus

83

— punjabensis
— ramosus .

IO, II

ii

, 00

var. pyrenaicus . . . .10

— rectus ...... 79

— rhenanus . . . . . 1 1 , 8 1

— roberti ...... 83

— savini ..... 40, 67

— sedgwicki ...... 40

— solilhacus . . . . . .83

— tetraceros . . . .88, 39*. 40*. 41*

— verticornis . . . 51, 53, 54, 61

— wallichi ..... 32-34
affinis ...... 32

— yarkandensis . . . . 32, 33
Chaetocarpus . . . . . .167

Chalk 5

CHANDLER, M. E. J. . 147-187, 291-314

Chasmatosaurus . . . . .322

Cheirolepidaceae .... 129, 130

Chillesford Clay ..... 5

Chimpanzee teeth . . . 141, 143, 144

China ...... 219, 220

Chromate staining . . 145, 252, 259, 277

Cladophlebis . . . . . .102

CLARINGBULL, G. F. 144, 225, 252, 257, 268-270
CLARK, W. E. LE GROS . 139-146, 234-242
Coelodonta antiquitatis . . 88

Collagen ...... 255

Colpodexylon . . . . . .216

Columbea (Araucaria) . 104, 109, 118, 131, 132
Cones (Araucarian) . . . .no

COOK, S. F. . . . 143, 146, 266, 285

Cordaites . . . . . 193

Cretaceous fruits . . . . .149

Cricotus ...... 364

CROFT, W. N. ..... 292

Cromer ...... 5

Croydon, Surrey ..... 307

Cryptomeria .... 124, 129, 130

Cuisian ..... 292, 294

Cupressaceae . . . . .122

Cycadales ..... 109, 121

Cyclopteris . . . . . .214

Cyclostigma . . . . . .198

— aegyptiaca . . . . .216

— kiltorkense . . . 199, 203, 206

— pacifica 202, 204, 206, 211, 212 ; PI. 17,
figs. 5-7 ; PI. 18, figs. 8-10 ; PI. 19, figs, ii,
13, 14 ; PI. 20, fig. 146 ; PI. 24, fig. 30.

— cf. — var. 212 ; PI. 23, fig. 290 ; PI. 24,
figs. 296, c

• — decorata . 206 ; PL 19, figs. 12, 14

— ungeri . . . . . .211

Cyclotosaurus ..... 385

Dama .

— clactoniana

— nestii

eurygonos

nestii .

Damacerus

— bessarabiae

ii, 90

81, 82*. 87, 90,91
86
82

13, 79, 80*. 81, 88, 90, 91
9-i I, 13
8, 9, 13, 90

— variabilis . . . . . 9, 13, 90
Danian . . . 150, 169, 175, 186, 187
Dano-Montian 149, 150, 159, 165, 167, 169,

178, 179, 183-186

DAVIDSON, C. F. . 225, 252, 262, 276-282

DAWSON, C. 144-146, 229, 230, 260, 268, 285
DE BEER, G. ..... 228

Dendrerpeton . . . 320, 882-384

Devonian ..... 320, 329

Dicerorhinus etruscus .... 86

Dicotyledones . . . . .165

Dicroceros . . . . . .13

Diospyros . . . . . .172

— schweinfurthi . . . 150, 168, 172

— sp. . 154*

Donetz Basin ..... 218

Dorset (localities) . . . 291, 297, 299
DOTHIE, H. J. . . . . . 251

Drillings, bone . . . 146, 255, 258
Dvinosaurus 321, 323, 329, 337, 342-344, 365,

366, 370, 383-385, 386*, 387*, 388

Ebenaceae
EDMUNDS, F. H.
Edops .
Edopsids
EDWARDS, W. N.
Egypt .
Elaphurus
Elaphus
Elatides

150, 172

225, 230, 232, 273-275

• 344. 358

• 343

. 141

149, 187, 203, 216, 217
ii, 14

35
129

Dadoxylon

1 06

Electron-microscope photographs 255 ; PL 30.
Elephas 246, 253, 263, 265, 269, 279-282, 284

— antiquus ..... 86, 87

— meridionalis . . . . 85, 86

— planifrons 85, 248-250, 263, 269, 279-281 ;
PL 30

— primigenius . . . . 88, 89

— trogontherii ..... 86

Elpistostege ...... 329

Encephalartos . . . . .109

Eoanthropus . . . . . .146

— dawsoni .... 230, 231, 233
Eobrachyops 329, 337, 343, 344, 363-366, 370,

377. 384. 385. 388

— casei .... 329, 356*, 384

— townendae . 329, 344, 346*. 347*. 348*,
349*. 353*. 354*. 356, 363*. 384, 386*. 387*.
388*

Eocene . . . . 159, 1 80, 187

— fruits ...... 149

Eoliths 243-245

Equus . . . . 253, 279, 280

Eremopteris . . . . . .195

— elegans ...... 195

— missouriensis . . . . 193

— peruianus . 192, 195, 197, 199, 202, 208, 212

— cf. sanjuanina . . . . 201, 214

— whitei . . 192, 195, 199, 201-203
Erpetosaurus . 329, 366, 870, 376*. 377, 384

396

INDEX

Erpetosaurus laevis
— radiatus

— tabulatus .
Eryops .
Esna Shales

184, 187
Estheria

Eucladoceros teguliensis
Euctenoceros .

— boulei

• 37°. 374*
37°. 372*. 373*. 375. 376*

367

344. 358, 359. 383. 385
159, 165, 167, 169, 176-179, 183,

IO2
40

ii, 14, 37, 45, 78, 90, 91

45

— ctenoides 37, 38, 40, 41*, 44*, 45, 46, 67, 73,
76, 78, 86, 88, 90, 91

- dicranios ... 38, 41, 45, 73, 86

— falconeri. . . .38, 43*. 45, 83, 85, 91

— incertae sedis . . 41*. 44*, 45, 46*

— sedgwicki 37, 38, 40, 42*, 43*, 44*. 45, 46,
53. 73. 76- 79, 83, 89-91

— sp. . . 44*, 46*

— teguliensis . . . 37, 38, 40, 91

- tetraceros 38, 39*. 40*, 41*. 45, 46, 76, 78, 86,
88, 90, 91

Eugyrinus .... 329, 882-384

— wildi ..... 320, 377
Euphorbiaceae . . 150, 166-168, 175
Euprox . . . . . 10, 13
Eutacta (Araucaria) . . . 104,132
Evotomys sp. ..... 89

151, 169, 175

150

180, 183

. 231, 243
143, 233, 248, 250, 256, 262,

Farafra, Egypt
Fayum, Egypt
Flacourtiaceae
Flint implements
Fluorine analysis

266

Fontechevade II skull section . . . 259*
Forest Bed, Norfolk . . . 3, 5. 84
FOSTER, A. . . . . . 144, 262

FRIEDERICHS, H. F. . . 232, 233, 285
FRYD, C.F.M. . 143, 225, 256, 262, 266, 267

Gazellospira torticornis . . .86

Gelatine . .;•••* . . .144
Geological Survey Museum . 4, 32, 54, 276
Germany . . . . 318, 329, 384

Gerrothorax 323, 329, 340, 341, 344, 384, 385

— pulcherrimus . . 341, 342*, 386*, 388*

— rhaeticus . . . . . .341

Gibraltar skull section .... 259*

Ginkgoales . . . . . .121

Gleichenia . . . . . 292

Glossopteris . . . • . ' . .322
Gnetales ...... 121

Gonioglyptus . . . . . .322

Government Chemist's Department 256, 262,

266

Gravel, Piltdown . 229, 243, 257, 268, 273

Greenland ..... 320, 329

GREGORY, J. W. . . . . 203

Gulo gulo ...... 86

GUNN, J. . . . . . . 3, 4

Gypsum in bones . 268-270

Hampshire (localities) . . . 291, 293
HASSAN, M. Y. . . . . 149, 185

Hausmannia . . . . . .102

Headon Beds ..... 292

HEIZER, R. F. . . . 143, 146, 266, 286

Helenia ...... 218

— bella 218

— inopinata ...... 218

— similis . . . . . .218

Heleniella theodori . . . . 218, 219

Heteroprox . . . . . .13

HEY, M. H. . . 145, 225, 252, 257, 268-270
Hippopotamus 86, 250-252, 263, 269, 279, 282

HOATHER, R. C. . " . . . .268

HOPWOOD, A. T. . 229, 230, 248, 251, 258

HOSKINS, C. R. . . 141, 146, 233, 283

HOWELLS, W. 233, 286

Hugonia . . . . . .174

Hura . . . . . . .175

Hydropotinae ..... 7

Icacinaceae . . 150, 152, 178, 179, 184

Icacinicarya .... 178, 179, 180

— platycarpa . . . . .179

— ? sp. 151, 179, 184; PI. 15, figs. 52-54;
PI. 1 6, fig. 66

— youssefi . 151, 178, 179 ; PI. 15, figs. 49-51
Ichthyostega ..... 320, 329
India . . . 318, 321, 322, 329, 330, 384
Intermedia (Araucaria) . . . 104, 132
Ipswich Museum . . . . . 4, 50
Isle of Wight .... 291,292

Joggins, South

JONGMANS, W. J.

• 377
189-224

KEITH, A. . . . . 229, 231, 235

KENNARD, A. S. . . . . . 246

Kharga oasis . 149, 169, 178, 183-185

Knorria ..... 191, 192

Kosseir, Red Sea 149, 159, 165, 167, 169, 178,
179, 184, 186

Labyrinthodonts . . . . .317
Lagenoidea ..... 152, 166

— bilocularis 151, 167, 168 ; PI. 12, figs. 21-23

— trilocularis 151, 167 ; PI. 12, figs. 16-20

La Plata Museum

LEAKEY, L. S. B. .

Lebachia

Lepidodendron acuminatum

— calamitoides

— caracubense

— cormgatum

— glincanum

— kidstoni

— leeianum .

— lissoni . . .

— nothum

— obovatum

— osbornei

HALL, E. T. .
Hailing skeleton
Hammerstones

145. 244
264

245- 246

200

233. 286
• 134
219
219

218, 22O
217, 219

2l8

219

219, 22O

198, 202, 207

218

192, 195, 197, 202, 206
219

— peruvianum 197, 199, 200, 202, 206, 207, 209

— rimosum . . 192, 195, 197, 202, 206

— robertii . . . . . .219

— spetsbergense . . . 198, 207, 219

INDEX

397

Lepidodendron sternbergi

— stylicum . . ,

— veltheimi . . ,

— veltheimianum .

— cf. volkmanni

— volkmannianum

. 191

. 218

192, 217

2OI

IQ2

217, 219

Lygodium smithianum
— volubile
LYNE, C. W. .
Lystrosaurus .

.311

308, 311

235. 239, 286

• 322

Lepidodendropsis 191, 198, 200, 206, 210, 211,
215, 216-220

— cyclostigmatoides .... 207

— de voogdi 209, 210, 216 ; PL 21 ; PL 22,
figs. 24-27 ; PL 25, fig. 32

— fenestrata .... 198, 207, 216

— hirmeri ..... 215—219

— lissoni . . . 207 ; PL 20, fig. 16

— mosaicum . . . . .217

— peruviana 198, 202, 206, 207, 209 ; PL 20,

fig- 15

— sinaica ... . 217

— sp 207 ; PL 20, fig. 17

— steinmanni 210, 211; PL 22, fig. 28a, d; PL 23,
fig. z8b, c; PL 24, fig. 31

— vandergrachti . . 210, 211, 215, 217, 218
Lepidophloios cf. laricinus . . .216
Lepidophyllum lanceolatum . . . 209
Lepidosigillaria . . . .217, 220

Lepidostrobus . 192, 196, 203, 206, 209

Lepospondyli ..... 329

Leptobus ...... 86

Leptophractus ..... 320

Libralces . . 15, 16, 19, 22, 29, 30, 86,90

— gallicus 7, 15, 16, 17*, 18*. 19, 20*, 22, 24,
25*, 26, 27, 28*, 29, 30, 31*, 78, 86, 88, 90

— incertae sedis .... 30, 31*

— latifrons . . 27, 28*, 29*. 30, 89, 90

— minor . . . 18*. 27, 28*. 89, 90

— reynoldsi 4, 19, 20*, 21*, 22, 23*. 24*. 25*.
26, 29, 30, 31*, 86, 88-90

Linaceae . . . . . .174

Linton, Ohio . . .319, 320, 375

Lloyds' Site, London . . 254, 255, 264
London Clay . . . . 149, 150, 152

Loxomma ...... 343

Loxommidae . 320, 329, 343, 383, 384

Lutetian .... 149, 292, 294

Lygodium ..... 304, 307

— articulatum . . . 308, 309, 311

— borneense . . . . . .311

— circinnatum .... 308, 311

— cubense . . . . . .311

— elmeri ..... 304, 311

— flexuosum ..... 308

— gaudini ...... 312

— hians . . . . . .311

— japonicum .... 308, 311

— kaulfussi 291-293, 307, 308, 309-312, 314 ;
PL 37 ; PL 38, figs. 84-87

— lucens . . . . . -3H

— merrilli . . . . . .311

— palmatum .... 308, 311

— polymorphum . . . . .311

— poolensis 291, 292, 312, 313, 314 ; PL 38,
figs. 88-96

— prestwichii 291, 293, 307, 308, 309 ; PL 36,
figs. 66-69

— reticulatum . . . 311, 313, 314

— salicifolium ..... 308

— scandens 311, 313, 314 ; PL 38, figs. 97-101

— skottsbergii . . . 308, 310, 312

Macacus ...... 86

Malta ..... 251, 265, 282

MARSTON, A. T. . . 232, 233, 258, 286
MARTIN, A. V. W. . . . . . 255

Mastodon arvernensis 85, 247, 250, 263, 269,

279-281
M astodonsaurus . . . . .318

— giganteus ...... 318

Megaceroides ..... 47, 48

Megaceros . n, 13, 46, 47, 48, 53, 85, 90, 91

— algericus .... 47, 51, 52

— antecedent .... 48, 50, 88

— belgrandi . . . . -51, 52, 67

— carnutorum ..... 45

— cazioti ..... 51.52

— cretensis . . . . . 51, 52

— dawkinsi 6, 49*, 51, 58*, 59*, 60, 61, 62*,
63*, 64*, 65*, 66*, 67, 68, 71*. 72, 89 — 91

— dupuisi .... 52, 53, 60, 69

— euryceros ..... 48, 52

— flabellatus . . . . 48, 51

— giganteus 48, 50, 53, 60, 62, 63, 66, 68, 72,
83, 91

— incertae sedis . . . . 71*, 72

— messinae ...... 53

— mosbachensis . . . 51, 52, 60, 66

— ordosianus .... 48, 51

- pachyosteus . . . 47, 48, 50-52

— pliotarandoides . . . . 51, 52

— savini 48, 49*, 50, 52, 58*, 59*, 62*, 67, 68*,
69*, 70*, 71*, 72, 83, 89, 90, 91

— solilhacus ..... 51, 52, 83

- verticornis 4, 48, 49*, 51, 52, 53, 55*, 56*.
57*, 58*, 59*. 60*. 61, 62, 63*, 68, 69, 71*,
72, 73, 86, 88, 90, 91

- yabei ..... 48, 51

Megalichthys . . . . . .318

Meganthereon meganthereon ... 86
Menispermaceae . . . . .184

Mesembrioxylon . . . . .106

Mesocalamites . . . . .197

Metbposaurs . . . . . .321

Micheevia ...... 218

— pulchella ...... 218

— rimnensis . . . . . .218

— uralica ...... 218

Microtus . . . . .86, 87, 90

— arvalinus ...... 89

— nivalinus ...... 89

— nivaloides ..... 89

— ratticeppoides ..... 89
MILLER, G. S. . . . 230, 232, 286
Mimomys ...... 86

— intermedius ..... 89

— maiori ...... 89

— pliocaenicus ..... 89

— savini ...... 89

Mindel-Riss interglacial . . . . 7, 34

Miocene ..... 13, 14

Mokattam, nr. Cairo . . . .159

Molar teeth, abrasion of . . . 142,236

398

INDEX

MONTAGU, M. F. ASHLEY
MORRIS, H. .
Moss, A. A. .
Muntiacinae .
Myrtaceae

Namurian
Nasal bones .
Natsiatum eocenicum
Nemorhoedus meneghinii .
Neorhachitome
Neurocardiopteris .
Newington, Kent .
New South Wales .
NEWTON, E. T.

229, 233, 286

• 244
145, 244, 245, 262

. 7. 10. i3
. 151

203
241, 258, 262

• 179

86

• 359
. 216

• 307
220
250

Nipa . . 149, 152, 153, 155, 162-164

— burtini 151, 159, 160-162 ; PI. 10, figs. 1-5 ;
PI. ii, figs. 7-9

— fruticans 160, 164; PL 10, fig. 6; PL n,
figs. 10-12

Nipaceae . . . . . .159

Nipadites sickenbergeri . 150, 159, 161, 162

Nitrogen content of bones — teeth 143, 254-256,

267
Norfolk ... . . . . 3

Norwich Museum 3, 19, 29, 42, 43, 49, 54-56,

58, 61, 67, 68, 72, 77, 83
Nova Scotia ..... 319, 377

Nummulites . . . . . .182

Nyctereutes megamastoides ... 86
Nymphaeopsis bachmanni . . . 151, 183

OAKLEY, K. P.

Odocoileinae .

Odocoileus

Oligocene

Ona Indian skull .

Orang utan .

— molars

Osteolepid fishes

Otocratia

Ouse 50 ft. terrace .

Ovibos moschatus

139-146, 243-265

7
15

. 151

258, 259

. 141

232, 234, 240, 241

- 319

• 329
273-275

Pagiophyllum . . . . .105

Palaeolithic implements 243, 244-246 ; PL 29
Palaeoplatyceros . . . . .13
Palaeowetherellia 150, 152, 159, 168, 171, 172,
i74. 183

— schweinfurthi 150, 151, 154*. 168, 174, 176 ;
PL 12, figs. 24-30; Pis. 13, 14; PL 15, fig.
48

Palmacites vimosus . . . .150

Palmatopteris furcata . 192, 194, 196, 202
Palm wood ..... 107, 133

Panchets ...... 322

Pandanaceae . . . . .151

Papaver ...... 182

Paracas flora ..... 208

Pararaucaria .... 119-135

— elongata ..... 119, 120

— patagonica 99, 119, 120, 123*. 125*, 127*.
128, 132, 134, 135 ; Pis. 6, 7

Paris Palaeontological Museum . . 52
PARSONS, L. E. . . . . 146, 326

• 97

• 259
366, 370, 377- 385

329, 367, 368*, 369, 370

• 329, 337

. 323, 338

216

Patagonia
Pathology of bone .
Pelion .

— lyelli
Pelorocephalus

— mendozensis
Pennsylvania

Peridermium galls . . . . .109
Permian 318-323, 328, 329, 342-344, 364, 365,

384

Peru . .... 189-224

Phyllospondyli ..... 320

Piltdown . . . 141, 229, 273, 274*

— canine .... 141, 271 ; PL 9

— gravel . . 229, 243, 257, 268, 273

— implements . . . 243 ; Pis. 28, 29

— mammalia ..... 247

— mandible 141, 235*, 254; Pis. 8, 27; PL 30,
fig. ii

— molars ..... 142, 256

— skull .... 141, 229, 259*
analysis .... 257, 262

— teeth 142-144, 234, 235*, 236*, 241*, 271* ;
Pis. 9, 27

Pinaceae . . . 121-124, I2^. 128-130
Pinakodendron . . . 194. 210, 215
Pinus ..... 106, 122-124
Pitimys arvaloides ..... 89

— gregaloides ..... 89
Plagiosauridae 321-323, 329, 340, 341, 342,

385, 388

Plagiosaurus depressus . . 321, 341, 384
Plagiosternum .... 323, 340

— granulosum . . . 321, 341, 384

— pulcherrimum . . . . 321, 341
Plagiosuchus . . 323, 329, 341, 384, 385
Platyceps . . . 323, 329, 343, 344

— wilkinsoni 322, 338, 339,* 384, 387* ; PL 39
Platystegos . . . 320, 329, 382-384

— loricatum . . . 377, 378*, 379*, 380*
Pleistocene . . . . . .11

PLESTERS, R. J. . . . 225, 271, 272

Pliocene . . . . 10, ii, 13, 14

Pliocervinae . . . . . 8, 10

Podocarpaceae . . . 122, 133, 134
Pontian . . . . 10, n, 13, 14

— deer 8-10

Porodendron . . . . . 217, 220

Portslade ...... 249

Potassium in fossil bones . . 260, 277

Proaraucaria . . . 109, 117, 118

— mirabilis .... 102, 104, no

— — var. elongata . . . .in

— patagonica . . . • no, in
Procamptoceras brivatense ... 86
Procapreolus . . . . . .10

Procervus ...... 9

— variabilis . . . • . . 8, 90
Prolepidodendron breviinternodium • .216
Protolepidodendron . . . .216

— ? arborescens ..... 220

— lineare ...... 220

— primaevum . . . . .220

— scharyanum . . . .216, 220

— yalwalense . . . . .220
Protolepidodendropsis frickei . . 218,219

— pulchra . . . . . .218

Przewalskium . . . . ii, 32

INDEX

399

Pseudo-A raucaria
Pseudovoltzia
Pteris prestwichi
Ptilophyllum .
PYCRAFT, W. P.

129
J33
3°7
1 02
231

Radioactivity of fossil bones 248, 249, 260, 276,

283, 284

Radiographs 142, 143, 146, 235, 241 ; PI. 9
RAMSTROM, M. ... 232

RANDALL, J. T. . . . . . 255

Rangifer arcticus ..... 88

— tarandus ...... 84

Rangiferinae . . . . . . 7

Reading Beds ..... 293

Red Crag . . 246, 248-250, 265, 280, 281
Redwood ...... 130

Reinschia ...... 327

Rhabdocarpus .... 192, 201

Rhachitomi ..... 320, 343

Rhacopteris . . . 195, 197, 200, 209, 216
• — asplenites . . . . . .195

— circularis 191, 197, 199-202, 208, 212, 213,
214, 216, 217 ; PL 25, fig. 38 ; PL 26, fig. 36

— cf. cuneata 199, 200, 202, 207 ; PL 20, fig. 18

— elegans . . . . . 195

— inaequilatera . . . . .192

— lindsaeformis . . . . .216

— ovata 199-202, 208, 209, 212, 213, 214, 217,
220 ; PL 25, figs. 33-37 ; PL 26, fig. 45

— peruianus

— petiolata .

— semicircularis

— transitionis
Rhaetic
Rhinoceros

— etrnscus
Rhodea

— filifera

— hochstetteri

— cf. moravica

— cf. stachei
ROMER, A. S.
Romeroites

— argentinensis
Royal Society
Royena

— desertorum
Rubiaceocarpum markgrafi
Rucervus

— eldi ....
Rusa ....
Russia

197, 202
. 197
201, 216

• 195
323. 329, 384

254, 255

247, 263, 269, 279, 280
193, 194, 196, 202, 216
192, 196, 202
195, 203, 204, 216
. 217
. 217

356, 369, 37°
129, 130

• 133
. 389
. 172

150, 168, 172, 176
151, 159, 163
ii, 14
. 15*
ii
318, 323, 329, 342, 384

SAINTY, J. E.

Salamandr aides giganteus

Saurerpeton .

SAVIN, A. C. .

Schizaeaceae .

Sciadopitys .

Scotland

SCOTT, D. B.

Securidaca tertiaria

Sedgwick Museum, Cambridge

Sequoia

— sempervirens

. 4, 7, 18, 29

. 318

321, 344, 366, 367

. 3, 4. 6. 7
291-314
130, 133

• 329
256

. 150

no

106, 122, 130

122, 130

. 3i8

• 343

• 259
229

. 329, 384

J93. 198, 202, 204, 205, 215

193, 217

218

Seymouria
Seymouriamorphs .
SHATTOCK, S. G.
Sheffield Park, Sussex
Siberia .
Sigillaria

— brardi

— canobiana

— fezzanensis . . . . .217

— mutans ..... 194, 205

— tessellata ...... 191

— tschirkovaeana . . . . .218

— youngiana . . . . .218

Sinai ...... 216, 217

Sinanthropus site ..... 47

Sinomegaceros .... 47, 48

Siwalik Beds . . 248, 249, 265, 280

SMITH, G. ELLIOT ..... 231

SMITH, R. A. . . . . . 246

South Africa (localities) 318, 321-323, 329, 331,

336, 384

South African Museum
Spain .
Sphenophyllum

— geigense

— saxifragaefolioides
Sphenopteridium . . . 193

— norbergii .....
Sphenopteris affinis . 192-194, 196

— asplenites .....

— bodenbenderi ....

— furcata .....

— hartlebeni . . . 191, 194

— paracasica . 194, 196, 197

— parasica

— whitei
Spitsbergen
STEEN, M.
Stegodon

Stegops .....

Stephanian .....
Stephanocemas ....
Stereospondyli ....
Stigmaria exigua ....

— ficoides .....

— stellata .....
Stigmariopsis ....
STRAUSS, F. B. .
Sublepidodendron antecedens

— fallax .....

— mirabile .....

— nordenskioldi ....

— subfallax .....
Sulphate in bones — teeth 252, 257, 259
Sus scrofa .....

— strozzii .....
Swanscombe Skull section

Sweden .... 329,

Swedenborgia ....

Syria ......

202, 203, 216 ; PL 17

218

366-371

247

• 336
. 217

• 193

216, 217

216, 217

-196, 216

193, 196

202, 203

• 195
196

192, 193
2O2, 2O3
199, 2O2
199, 2O2

, figs. 1-4

336, 384

, 374-376

248. 253

376, 377

• 194

13

343. 357
219

. 217

. 199

251, 262

. 218

. 219

219

219

219

266, 268
. 86
. 86

• 259*
341- 384

• 133
. 216

Taiwania . . . 126, 129, 130, 134

Tapirus arvernensis . . . .85

Taxaceae . . . . . .122

Taxales 134

Taxodiaceae 99, 118, 122, 124, 126, 128-130,
132-135

400

INDEX

Taxodium

Teichosperma spadiciflorum

TEILHARD DE CHARDIN, P.

Tethyan flora

Tethys ocean .

Texas .

Thiebaudia .

122, 126, 130
. 151

152-158

152, 153. 155, 156, 158
• 329, 344. 362, 364, 384
180

— rayaniensis 151, 159, 180, 183 ; PI. 16, figs.
58-63

THORVILDSEN, K. . . . . . 255

Tilbury (Kent) skull . . . .264

Tollund Man 255

TOOMBS, H. A. . . . . 229

Trachyphyton neglegibile . . 199, 203, 206
Trematosaurs . . . . -336
Trias 318, 319, 321-323, 329, 330, 342, 343,

357- 384

Trimerorhachis 321, 329, 344, 355-357, 358, 359,
363*-366, 375

— conangulus ..... 363

— insignis . . . . . 363

— sp. . . . 358*, 360*, 362*. 363*
Trimerorhachoidea .... 343

Triphyllopteris

— collombiana
figs. 40-42

— lescuriana

— minor

— ? peruviana

201, 209, 214, 216
199, 203, 214, 217 ; PI. 26,

214, 215 ; PI. 26, figs. 43-45
. 217

215 ; PI. 26, fig. 46
Trochophyllum breviinternodium . . 216
Trogontherium boisvilletti ... 86

Trucheosaurus . . 327, 329, 330, 384

— major 327, 328*

Tungussogyrinus ..... 329
Tunisian fossil site . . . 249, 280

Turbinal bone . . . 241, 258, 262

Ural Mts

Uranium in fossil bone .

Ursus arvernensis .

Vandyke brown on Piltdown canine

Vertebraria ....

Villafranchian

Voles .....

Voltzia .

. 218
248, 276, 283

• 249

257, 272

• 322
230, 247, 248

86, 89

• 133

Wadi Rayan, Egypt
Wales ....
WATERSTON, D.
WATSON, D. M. S. .
WEIDENREICH, F. .
WEILER, G. .
WEINER, J. S.
WEINERT, H.
WELLS, R. A.
WERNER, A. E. A.
Westphalian .
Wetherellia
— dixoni
— variabilis .
Weybourn Crag,

WrISEMAN, J. D. H.

Wood, coniferous 105, 134,

WOODHEAD, S. A. .

WOODWARD, A. S. 144-146,

253
Woolwich & Reading Beds

149, 180
216

230, 287

315-392

231-233, 287

251, 262, 267

139-146, 229-233

231, 233, 287

• 283
271, 272
193, 203

152, 171, 172, 174
150, 154*. J?2-i74

150, 154*, 172, 173

3

. 144
; PI. i, figs. 2, 3, 7

• 257
229-232, 242, 243,

293, 307-309

X-ray crystallographic analysis 257, 268 ; PI. 31
— spectographic analysis . . .244

Ularia ovalis
Ullmannia
UNDERVVOOD, A. S.
University College, London

219

• 134

235, 240, 287
. 389

YOUSSEF, M. I. . . 149, 176, 186

^atrachys ...... 376

Zoological Museum, Cambridge . . 342

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