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ANNALS: 


OF THE 


SOUTH AFRICAN MUSEUM 


VOLUME XXXI 


vv ee ee 2 ee rey =e TE —— | ee es eer ers 


ANNALS 


OF THE 


SOUTH AFRICAN MUSEUM 


VOLUME XXXII 


PRINTED FOR THE 


TRUSTEES OF THE SOUTH AFRICAN MUSEUM 


AND THE 
GEOLOGICAL SURVEY OF SOUTH AFRICA 
BY NEILL AND CO. LTD., 212 CAUSEWAYSIDE, EDINBURGH. 


1934-1950. 


ao]. be 


TRUSTEES OF THE SOUTH AFRICAN MUSEUM. 


Professor R. S. Apamson, F.R.S.S.Afr. 
The Rev. S. J. Fort. 

Professor D. L. ScuHoutz, D.Sc. 

C. J. SIBBETT, J.P. 

S. H. Sxaire, Ph.D., F.R.S.S.Afr., J.P. 


SCIENTIFIC STAFF OF THE SOUTH AFRICAN 
MUSEUM. 


KepreL Harcourt BaRNnarD, M.A., D.Sc., F.L.S., Director; in Charge of Fish 
and Marine Invertebrates. 

ALBERT JOHN Hesse, B.Sc., Ph.D., F.R.E.S., Assistant in Charge of the Ento- 
mological Department. 

Miss G. Joyce Lewis, B.A., Assistant in Charge of the Botanical Department. 

LizUWE Dirk Boonstra, D.Sc., Assistant in Palaeontology. 

A. J. H. Goopwin, M.A., Honorary Keeper of the Ethnological and Archaeological 


Collections. 
Miss K. Marcaret Suaw, B.A., Assistant in Charge of the Ethnological Collections. 


LIST OF CONTRIBUTORS. 


R. S. ADAMSON. 
Fossil Plants from Fort Grey, near East London 


L. D. BoonstTRa. 


Pareiasaurian Studies. The Cranial Date eer 

The Dermal Armour : 

The Vertebral Column and Ribs 

A Contribution to the Morphology of the Gorgonopsia : 

Additions to our Knowledge of the South African Gorgonopsia preserved 
in the British Museum : 

A Contribution to the ee ee of the Mammal-like Reptiles of the 
Suborder Therocephalia_ . 


L. CAYEUX. 
The Phosphatic Nodules of the Agulhas Bank 


H. B. 8. Cooke. 
A Critical Revision of the peli aay baeisaact of Southern 
Africa : ; : : 


S. H. Haveuton. 
On some Karroo Fishes from Central Africa 
See also J. V. L. Rennie 
J. V. L. RENNIE. 
On Placocystella, a New Genus of Cystids from the Lower Devonian of 
South Africa ' : 
Lower Cretaceous Lamellibranchia from Northern Zululand (with an 
Account of the Geology of the Cretaceous Beds and a Preliminary 
Analysis of the Associated Ammonite Fauna by S. H. Haughton) . 


LIST OF NEW GENERA AND SUBGENERA 
PROPOSED IN THIS VOLUME. 


Aelurosauroides (Gorgonopsia, Reptilia) Boonstra 
Arctognathoides (Gorgonopsia, Reptilia) Boonstra 
Ischnolepis (Pisces) Haughton . 

Megacucullaea subg. (Arcidae, Mollusca) Ronnie 
Placocystella (Cystoidea) Rennie ; 
Sphenotrigonia subg. (Trigoniidae, M oluxea) Rennie : 


393 


97 


269 


277 


190 
143 

97 
305 
269 
365 


PLATES 


XVIII 


XXX. 
XXXI. 


XXXII 


XXXII. 
XXXIV. 

XXXV. 
XXXVI. 


XXXVII 
—LY. 


LIST OF PLATES. 


Pareiasaurian skulls. 


Pareiasaur teeth. 

Bradysaurus baini. 

Bradysaurus seeleyt. 

Bradysaurus seeleyt and vanderbyli. 
Bradysaurus vanderbyli. 
Bradysuchus whaitsi. 
Nochelesaurus alexandert. 
Nochelesaurus strubent. 
Nochelesaurus strubent and Dolichopareia angusta. 
Dolichopareia angusta. 
Brachyparewa rogersi. 

Brachypareia watsont. 
Embrithosaurus schwarz. 


XIX. \ Pareiasuchus peringueyt. 


XX. 
XXI. 
XXII. 
XXIII. 
XXIV. 
XXYV. 
XXVI. 
XXVII. 
XXVIII. 
XXIX. 


Pareiasuchus nasicornis. 
Pareiasuchus nasicornis and Anthodon serrarius. 
Anthodon serrarius and Pareiasurus serridens. 


Dermal Armour of Pareiasaurians. 


Bradysaurus seeleyt. 

Fossil Karroo Fishes. 

Map of Agulhas Bank Area. 
Phosphatic Nodules. 


Devonian Cystids. 


Lower Cretaceous Lamellibranchia. 


DATE OF ISSUE OF THE PARTS. 


Part 1, pages 1-136, April 1934. 

Part 2, pages 137-268, July 1934. 

Part 3, pages 269-392, August 1936. 
Part 4, pages 393-480, December 1950. 


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INDEX OF GENERA AND SUBGENERA. 


PAGE 
A 
Aelurognathus . é : 2 143 
Aelurosauroides ’ : ye Le) 
Aelurosaurus . ; : 2 ile’) 
Alectryonia : : ; -al6 
Alopecodon : : B +. 216 
Anthodon t 35, 42, 65 
Arctognathoides : : . 148 
Arctognathus 142, 203 
Arctops : : 5 se dls) 
Asinus . : : : 451 
Atherstonia : 3 ; lo? 
Aucellina . : : ‘ «ol 
Avellana . : ‘ : . 388 
B 
Brachypareia 31, 41, 64 
Bradysaurus 24, 39, 63 
Bradysuchus . , ‘ Sea 
C 
Cardium ...-> *. : : i 382 
Ceratotherium . : ; . 404 
Cerdodon . 4 ; : : ez 
Cerdognathus . ; : . 211 
Cucullaea . ; ; 3 . 304 
Curtisia . : : : 2388 
Cynariognathus ‘ : . 216 
Cynariops ‘ : i = Lo 
Cyniscodon : ; : ee NILES) 
Cynodraco 158, 210 
Cyphoxis : : ; . 3804 
D 
Diceros : : ; : . 401 
Dicroloma ‘ , . 388 
Dolichopareia 


1x 


EB 


Elginia . : 
Embrithosaurus 
Equus 
Euchambersia 
Kurygnathohippus 
Exogyra 


Gervillia 
Glycymeris 
Goniomya 
Gorgonops 


Hipposaurus 
Hyorhynchus 


Icticephalus 
Ictidosuchoides 
Idonearca 
Inoceramus 
lotrigonia 
Ischnolepis 
Isognomon 


Koalemasaurus . 
Kolpohippus 
Kraterohippus 


Leptotrachelus . 


29, 64 | Lopha 


PAGE 


BF red, 
29, 42, 63 
431 

261 

423 

316 


310 
310 
378 
18] 


154, 158 
231 


186 
316 


x Index of Genera and Subgenera. 


PAGE 
Lycaenodon  . : : . 192 | Ptychomya 
Lycaenodontoides . : . 141 | Pygopterus 
Lycaenops : , : . Ld4 
Lycideops : : ; . 235 
Lycosaurus - . : . 202 
Rhinoceros 
M Rutitrigonia 
Megacucullaea . é : . 305 
Megatrigonia . ; : - idol 
Metaschizotherium . : ss g ed 
Moschorhinus . : : - 238 | Scabrotrigonia . 
Scutosaurus 
N Scylacops 
i Scylacosaurus 
Neithea . : ; . <a. O10 Be ee 
Nochelesaurus . : : 27, 63 Sipe eas . 
Notohipparion . e ’ -. 425 ee 
lee Steinmanella 
Notosollasia . : 2 . 242 St 
errohippus 
Stylohipparion . 
O 
Opsiceros . : : . 401, 404 
Theriognathus . 
P Thetironia 
Tigrisuchus 
Panopaea . ‘ é 5 . 385 | Transitrigonia 
Panope . : : : . 385 | Trigonarca - 
Panopea . : : : . 385 | Trigonia 
Paraeiasaurus . : . 34, 43, 65 | Trochosaurus 
Pareiasuchus  . : . 33, 44, 65 | Turritella . 
Pholadomya . : : ~ oe 
Placocystella. 5 : . 269 
Placocystis : ‘ : 7 209 
Platyoropha : : : . 24 
Pleurotrigonia . : . 361 | Veniella 
Podocarpus : é : ~ ip 
Pristerognathus : é a 2a 
Propappus : , . 32, 44, 64 
Protocardia : : ; a9 
Pseudavicula . ‘ : . 315 | Whaitsia . 


Pterodonta . .  .  . 388 | Widdringtonia . 


PAGE 
383 
101 


401, 404 
355 


378 


252 
75 


ANNALS 


OF THE. 


SOUTH AFRICAN MUSEUM 


LO: AXA. 


DESCRIPTIONS OF THE PALAEONTOLOGICAL MAPERIAL j 5) 
COLLECTED BY THE SOUTH AFRICAN Vie 


PART I, contarming :— | Yen ne aye 
i : : i ON “INA MU cs 
1. Pareiasaurian Studies. . = 


Part [X.—The Cramal Osteology.—By Lizruwe D. BoonstrRa, 
D.Sc., Department of Palaeontology, South African 
Museum. (With Plates I-X XII and Text-figures 1-5.) 


Part X.—The Dermal Armour. By Lizuwe D. Boonstra, 
D.Se., Department of Palaeontology, South African 
Museum. (With Plates XXIII-X XVII.) 


Part X1.—The Vertebral Column and Ribs. By Lizuwz D. 
BoonstrA, D.Sc., Department of Palaeontology, South 
African» Museum. (With Plate XXVIII and Text- 
figures 1-7.) 


a Fossil Plants from Fort Grey near East London. By Prof. 
R. 8. Adamson, M.A., Botanical Department, University 
of Cape Town. (With 27 Text-figures.) 


3. On some Karroo Fishes from Central Africa. By 8. H. 
HavueutTon, B.A., D.Sc., Hon. Keeper of Palaeontological 
Collections. (With .Plates XXIX-XXXI and 2 Text- 

_” figures.) 

A. The Phosphatic Nodules of the Agulhas Bank. (A Study of 
Submarine Geology.) By L. Cavzux, Membre de |’Institut, 
Professor at the Collége de France, Paris. (Translated into 
English by Dr. 8. H. Haughton.) (With Plates XX XII- 
XXXYV.) 


ISSUED APRIL 1934. PRICE 20s. 


PRINTED FOR THE 


~ TRUSTEES OF THE SOUTH AFRICAN MUSEUM 


BY NEILL AND CO., LTD., 212 CAUSEWAYSIDE, EDINBURGH. 


<a 


ANNALS 


OF THE 


SOUTH AFRICAN MUSEUM 


VOLUME XXXII. 


1. Pareiasaurian Studies. 


Part [X.—The Cramal Osteology.—By Lizuwe D. Boonstra, D.S8c., 
Department of Palaeontology, South African Museum. 


(With Plates I-X XII and Text-figures 1-5.) 


INTRODUCTION. 


In this paper the skull features are examined more fully than was 
possible at the time of the publication of “An Attempt at a Classifica- 
tion of the Pareiasauria based on Skull Features.”” An examination of 
the systematic aspect of the cranial features will now better serve to 
augment the conclusions arrived at in the series of papers which have 
dealt with the postcranial skeleton, where the tentative preliminary 
classification has been tested and found to hold good in all but one 
case, viz. Platyoropha broom. From these papers it has become 
obvious that the genus Platyoropha is invalid. The single specimen 
referred to it is in reality a member of the genus Bradysaurus. 

In the preparation of this paper I am greatly indebted to Dr. 8. H. 
Haughton, who has always been ready to offer help by discussion 
and criticism. I have also had access to his sketches and notes of the 
material in Kuropean institutions. To Mr. J. Hewitt, Director of the 
Albany Museum, I am grateful for the opportunity of examining the 
type skull of Koalemasaurus acutirostris. Through the kindness of 
Miss Wilman, Curator of the McGregor Museum, I have been able 
to study the Pareiasaur material in her charge. To Mr. Barnum 
Brown and Dr. W. K. Gregory, of the American Museum, I am 
indebted for the photographs of the Pareiasaurs in that institution. 

VOL. XXXI, PART 1. I 


2 Annals of the South African Museum. 


HISTORICAL. 


In his “Descriptive and Illustrated Catalogue of the Fossil Reptilia 
of South Africa in the Collection of the British Museum,’ Owen in 
1876 described three Pareiasaurs, viz. Parevasaurus serridens, Pareia- 
saurus bombidens, and Anthodon serrarius. Notwithstanding the 
fact that he had three skulls, he did not notice their stegocrotaphic 
nature, and he classified these three animals as zygocrotaphic Dino- 
saurs. The description (pp. 6-8) and two of the illustrations (pls. vi, 
vil) of Pareiasaurus serridens are based on a cast, the original being 
lost. Parevasaurus bombidens, described (p. 9) and figured (pls. vii, 
1X), Was a species founded on the snout of an animal. Owen confused 
the mandible with the maxillary portion of the skull, and vice versa. 
The specific differences in his two specimens rested solely on the 
nature of the teeth. In P. serridens there are 15-16 serrate teeth in 
both upper and lower jaws; but on the next page he says that 
originally only 14 teeth were counted. In P. bombidens there were 
19 teeth, which differ in form from those of P. serridens, viz. the 
outer surface of the maxillary teeth are more convex or bulging, the 
serrations are broader, and some of the linear intervals extend nearer 
to the base of the crown. Anthodon serrarius, described (pp. 14-15) 
and figured (pl. xii), differs, according to Owen, from the two species 
of the genus Pareiasaurus “in having the crowns more compressed 
transversely, and more expanded antero-posteriorly beyond the 
roots, which are relatively longer and more deeply implanted into the 
jaw; the crown terminates in a semicircular border, deeply notched, | 
rather crenate than serrate. . . .” 

In 1888 Seeley described (pp. 60-77) and figured (pls. xii—xv) a 
skull of a Pareiasaur from Palmiet Fontein. On plate xvi he also 
gives the palatal view of the cast of Pareiasaurus serridens. Although 
Owen had based his classification on the teeth, Seeley maintains that 
this is impossible, and says, “It seems to me that their differentiation 
must rest upon the form, proportions, and structure of the skulls.” 
But a few lines further on he acknowledges that “As very little is 
preserved of the skull of P. bombidens, and P. serridens is only known 
from the cast of the distorted skull and fragment of the lower jaw, 
no detailed comparison can be profitably pursued.”’ He then differen- 
tiates between P. serridens and P. bombidens, viz. the former is high 
and narrow, and the latter depressed and broad. As the specimen 
from Palmiet Fontein is also depressed and broad it is referred to 
P. bombidens. 


Pareiasaurian Studies.—Part IX. 3 


In 1892 Seeley obtained another specimen from Tamboer Fontein 
(skull described, pp. 315-320, figured pl. xx) which he also refers to 
P. bombidens Owen. A third specimen (skull described, pp. 322-329, 
figured pls. xvul, xvii, xix) from De Bad was described by Seeley as 
a new species—P. bain. The specific differences being “broader 
and relatively shorter, the rows of palatine teeth are closer together, 
16 teeth more slender, smaller, less inflated transversely, overlapping 
each other less, with the terminal denticles of the crown less pro- 
longed, choana larger, and more elongated.” 

Before considering the other Pareiasaurs it will be more profitable 
to see in what way attempts have been made to settle the taxonomy 
of these four Pareiasaur specimens. In 1914 Watson defined a 
number of Pareiasaurian genera. The two of interest here are: 
Parevasaurus—skull with deep cheek and pointed form, the type 
skull being the cast made by Owen; Bradysaurus—skull depressed 
and rounded, the type being the De Bad specimen, Br. baint. No indi- 
cation is here given to which genus Owen’s specimen and Seeley’s Pal- 
miet and Tamboer specimens are to be referred; but in a previous 
paper Watson states that P. serridens almost certainly is not co- 
generic with P. bombidens. It would therefore appear that Watson 
intended the three specimens of P. bombidens to be referred to the 
genus Bradysaurus. In their preliminary classification Haughton and 
Boonstra suggested that Owen’s P. bombidens be considered incertae 
sedis, and they proposed to raise Seeley’s two specimens to the rank 
of a new species—Bradysaurus seeleyi. As Owen's P. bombidens 
certainly cannot be a Pareiasaurus, and probably is a Bradysaurus, 
it appears advisable to refer to it as Bradysaurus ? bombidens. 

In 1930 Boonstra described a skull in the collection of the Univer- 
sity of Stellenbosch. This was identified as Pareiasaurus serridens, 
and as the type is lost this specimen was proposed as the neo-type. 
We thus have the following solution :— 


Pareasaurus serridens type skull: Owen’s cast in British Museum. 
Neo-type: skull in University of Stellenbosch. 

Bradysaurus baini type skull: Seeley’s De Bad specimen in British 
Museum. 

Bradysaurus seeleyi type skull: Seeley’s Tamboer specimen in 
British Museum. 

Bradysaurus ? bombidens type skull: Owen’s imperfect specimen in 
British Museum. 


In 1903 Broom (pp. 125-127, pl. xvi) described a nearly complete 


4 Annals of the South African Museum. 


specimen as Parevasaurus serridens. Watson subsequently showed 
that it cannot be referred to this species, and proposed the name 
Embrithosaurus schwarz. Of this specimen Broom described the 
teeth, and of the bosses on the skull he gave a detailed account. 

In 1910 Broom gave a restored figure of the palate of Pareiasaurus 
(fig. 1) with hypothetical sutures between the various palatal bones 
and an erroneous suture between the pterygoid and basioccipital. 
He also mentions (p. 201) a distinct septomaxillary. 

In 1912 Versluys gave a good figure of the palate of Parevasaurus 
baint without attempting to indicate the palatal sutures (fig. R). 
He also points out Broom’s incorrect determination of the pterygoid- 
basioccipital suture (pp. 364 and 613). 

In 1913 Broom and Haughton described (pp. 17-19, pl. v) the skull 
of a new Pareiasaur, which they named Pareiasuchus peringueyt. 
According to them it differs from the other Pareiasaurs in that the 
snout is nearly twice as broad as deep, the great size of the temporal 
roof and the fact that it 1s turned forwards with a huge rounded 
boss at the angle, the teeth are relatively larger and flatter than in 
Parevasaurus, and the cusps are either smaller or more rapidly worn 
down—there are 13 teeth in the upper jaw with 13-15 cusps. In 
the same year Broom gave a short account without figures of an 
imperfect skull, which he called Parevasaurus acutirostris. It differed 
from the then known Pareiasaurs in being two-thirds their size, and 
in a few other features. 

In 1914 there appeared a photograph, with a short caption, of a 
skull of a new Pareiasaur, Pareiasaurus whaitsi, in the American 
Museum Journal. In the following year the same illustration is 
given (fig. 1) accompanied by a short description. It differs from 
P. serridens and Br. baini in the much larger size of the orbit and in 
the lower jaw having two horn-like bosses instead of only one. 

In 1914 Watson gave an excellent account of the structure of the 
Pareiasaurian skull. This was the first account in which an attempt 
was made to consider the subject morphologically. Previous authors 
practically confined themselves to vague descriptions of shape and 
size, with haphazard identification of some of the constituent bones. 
Watson gives a sketch of the side of the partial skull of Anthodon 
serrarius and the palate of Bradysaurus seeleyi with the bones identi- 
fied, and a good account of the whole skull of a specimen from Hotten- 
tots River which he regarded as belonging to Embrithosaurus schwarz. 
In addition a detailed comparative account is given. 

In 1924 Broom described (pp. 499-504, figs. 1-2) the roof bones 


Pareiasaurian Studies.—Part IX. 5 


of a young skull of a Pareiasaur. In his determinations of some of 
the bones he rightly differs somewhat from Watson, and, furthermore, 
reluctantly accepts Watson’s classification. The lower jaw and 
teeth of a new form, Pareiasaurus strubent, is described and figured 
(pp. 507-508, figs. 5, 6). 

Such was the condition when Haughton and Boonstra presented 
their “Attempt at a Classification of the Pareiasauria based on Skull 
Features.’ Here a key was drawn up of 12 genera and 18 species, 
the basis of the classification being the nature of the teeth, the pro- 
portions of the skull, size, and ornamentation. This was followed by 
an account of the brain-cases of five species wherein attention is drawn 
to some specific differences. 

In the Annals of the University of Stellenbosch, Boonstra described 
the skull and brain-case of Pareiasaurus serridens, and showed that 
it differed from that of the other Pareiasaurs. 

In 1930 Broom published a note, with two figures, on what he 
considered to be a new species of Anthodon. 

Outside South Africa other Pareiasaurs had meanwhile also become 
known. Newton had already, in 1893, described Elginia mirabilis 
from the Elgin Sandstones in Scotland (pls. xxxvii—xl), and shown 
that this bizarre skull belonged to an animal related to the Pareiasaurs. 

Amalitzky, during his work on the North Dwina, had collected 
numerous remains of Pareiasaurs but never described them, although 
specimens were mounted and plates prepared. In 1922 Karpinsky 
published extracts of Amalitzky’s memoir. Here four species of 
Parewwsaurus (sic) were named, accompanied by very vague short 
descriptions. There are P. karpinskiw, P. horridus, P. elegans, and 
P. tuberculatus. Sushkin in his study “‘On the Modifications of 
the Mandibular and Hyoid Arches and their Relations to the Brain- 
case in Harly Tetrapoda” described and figured (figs. 23-26) the 
features relevant to his subject. The two chief points mentioned 
being the tubercle on the quadrate for the attachment of the 
stapes, and the fact that he maintains to have identified a supra- 
temporal bone in the “cheek.” 

Hartmann- Weinberg in 1930 gave a fairly full account of the skull 
of a North Dwina form, and establishes the fact long suspected by other 
authors that the Russian specimen cannot belong to the genus 
Pareiasaurus. For the specimen newly mounted by herself she 
proposes the name Scutosaurus karpinsky (karpinskuw ?). 

In 1931 von Huene, in his “Beitrag zur Kenntnis der Fauna der 
Siidafrikanische Karrooformation,”’ published some notes on the 
skull of Bradysaurus baini (pl. i, figs. 1, 2; pl. u, fig. 1). 


6 Annals of the South African Museum. 


GENERAL OSTEOLOGY. 


GENERAL FEATURES. 


The Pareiasaur skull is, as is the rule in the Stegocephs, in many 
cases broad and fairly depressed; but in some forms it is narrow 
and high. The temporal roof, although stegocrotaphic, is no longer 
primitive as it has lost one of the three original temporal bones. 
The quadrate retains the primitive rigid condition in being immovably 
attached to the ventral surface of the two temporal bones, and was 
apparently also cartilaginously attached to the brain-case. In 
addition the pterygoid is also immovably connected to the basi- 
cranium; this is, however, no primitive condition judging by the 
condition manifested in Crossopterygii, Stegocephalia, and primitive 
and other Cotylosauria; the Pareiasaur skull is therefore primarily 
monimostyle, but secondarily akinetic. Neither is the Pareiasaur 
skull divisible into two segments—mazxillary and occipital—movable 
relative to each other, the so-called metakinetic condition which 
still obtains in some Cotylosauria, e.g. Labidosaurus. The otic 
notch, well developed in Stegocephs and primitive Cotylosaurs 
(Seymouria), is practically obliterated. The primitive long and broad 
fenestra palatinalis or interpterygoid slit is, in the Pareiasaurs, 
secondarily partially or totally closed. The choanae retain their 
primitive condition far forward but are not confluent with each 
other. All skulls display a certain amount of sculpturing of the 
superficial dermal bones. In the smoothest skulls each bone has a 
small low central boss from which indistinct rugae and the fibres of 
the bone radiate; in addition, the posterior edge and angle of the 
“cheek” carry large, but low, bosses. In the more highly ornamented 
skulls the same ground plan is retained, but all the features are 
remarkably accentuated, so that in some cases actual horns are 
produced (compare the living forms Moloch and Phrynosoma); the 
bosses of the “cheek” are also increased in number and size. 

In dorsal view (PI. II) the Pareiasaur skull appears roughly tri- 
angular or in some cases quadrangular. The central part forms a 
fairly flat or slightly arched table bounded laterally by a ridge run- 
ning from the orbit to the tabular boss and from the orbit to the 
lateral margin of the nostril. This ridge is low in some forms but 
very strong in others. In addition there is in some forms a lateral 
bulging of the maxilla and lacrymal. It is thus clear that a central 
table is to be distinguished from the sides which is chiefly formed 


Pareiasaurian Studies.—Part IX. i‘ 


by the so-called bony “‘cheeks.”’ These “cheeks” lie in some cases 
nearly vertically down from the table; in others they are flattened 
out and diverge far laterally. The angle of the “cheek” lies far 
forward—up to the plane of the orbits—or nearly in line with the 
posterior border of the skull. The central table is generally more or 
less triangular in shape, the base formed by the posterior border of 
the skull, the apex by the internasal processes of the nasals; in some 
cases it is, however, nearly rectangular. This is due to the relative 
widths between the tabular bosses and orbits, and also to the direction 
of the ridge running from the orbit to the nostrils. The relation 
between the length and breadth of the dorsal table produces different 
types of skull, viz. long and narrow, or short and broad skulls. The 
posterior edge formed by the tabulars and postparietal is in most 
forms sharp and abrupt, but in some is bent down to form a flange 
partly overhanging the occiput. In dorsal view the orbits appear 
elliptical, and lie either mostly in the anterior half of the skull or 
mostly in the posterior half. The nostrils can generally be seen in 
dorsal view, although they must be described as more terminal than 
dorsal; in some cases they are overhung by nasal bosses; they vary 
in shape; in some cases the floor of the nostril and even the internal 
nares can be seen, in others not. 

In lateral view (PI. I) the Pareiasaur skull is roughly triangular in 
shape, the base being the posterior border of the “cheek ”’ and the 
apex the snout. In this view the rugosities on the cheek of some forms 
are well seen. Some skulls are fairly smooth, but even in them the 
structure and radiations of the bone-fibres show that all skulls are 
based on the same ground plan and that if bosses are not developed 
it just means that a potentiality has not been realised. Taking the 
family as a whole, the orbit can be described as more or less circular, 
although in some forms it is elliptical with the long axis lying longi- 
tudinally or obliquely. The nostril appears more or less kidney- 
shaped; in some forms it is partially obscured by the bulging of the 
lacrymal and maxilla. The height of the ‘‘cheek” is seen to differ 
considerably amongst the various forms, and this difference is mainly 
due to the size of one element—the quadratojugal. It also becomes 
apparent that amongst the Pareiasaurs there are high and depressed 
skulls; forms with more or less vertical “‘cheeks,” and forms with 
“cheeks ” diverging laterally. The tabular boss generally forms a 
prominent dorso-posterior angle, but in forms like Scutosaurus, 
Pareiasaurus, Anthodon, and Pareiasuchus the boss lies more on the 
lateral surface, and forms a much less prominent feature. From the 


8 Annals of the South African Museum. 


figures given in lateral view it is clear that the maxilla is not such a 
very low bone as Watson has maintained. 

In occipital view (Pl. IV) the Pareiasaur skull presents either a 
rounded arch, in the forms where the tabular bosses are not prominent, 
or a truncated V. The differences in height, the inclination and the 
rugosity of the “cheek” are very apparent. The shape and size of 
the post-temporal fenestra also varies considerably, this being 
largely due to the structure of the paroccipital process. The pos- 
terior edge of the tabulars generally forms a slight downhanging 
shelf, but the postparietal edge is horizontal; in Pareiasaurus serrt- 
dens both these elements form a large overhanging flange. Morpho- 
logically the most important feature is the strength of the supra- 
occipital pillar and its firm attachment to the postparietal. Both 
the squamosal and the quadratojugal posteriorly have supporting 
flanges for the quadrate, which stands either vertically or is inclined 
slightly in anterior direction. 

In ventral view (Pl. III and text-figs. 2 and 3) the primary palate, 
its dentigerous nature, the premaxillary processes and maxillary 
flanges, the forward position of the choanae, the immovability of the 
basisphenoid-pterygoid junction, the general absence (with a few 
exceptions) of an interpterygoid slit, the forward position of the 
inclined quadrates, are prominent features. Differences within the 
family—the nature of the median pterygoid suture, the inclination 
of the quadrate, the shape of the basisphenoid, the overhang of the 
maxillaries and premaxillaries, the shape of the choanae, etc.—are 
apparent from the figures. 


Nasal.—Together the two nasals form a rough pentagonal. An- | 
teriorly they clasp the lateral edges of the ascending premaxillary 
processes. They form most of the dorsal and median edges of the 
nostrils. As a rule each nasal carries a low boss from which the 
bone-fibres, and sometimes rugae, radiate to the edges of the bone; 
this boss is sometimes (Pareiasuchus nasicornis) forwardly directed 
and begins to look like a horn. In one form (Bradysuchus whaitst) 
there is, in addition to these two nasal bosses, one central boss of 
which half is contributed by each nasal; in another form (Nochele- 
saurus alexandert) this third boss lies at the junction of the nasal- 
frontal sutures and all four bones contribute towards it; in still 
another case (Nochelesaurus strubenz) there is only a single median 
nasal boss—half formed by each nasal. 

Frontal.—The two frontals together form a six-sided figure. The 


Pareiasaurian Studies.—Part IX. 9 


nasal and parietal sutures are parallel, whereas laterally the frontal 
edge is V-shaped. Roughly at the point of the V the pre-postfrontal 
suture enters. Hach frontal carries a boss from which the bone- 
fibres or rugae radiate; this boss is often situated in the V and 
sometimes coalesces with the pre- and postfrontal bosses; in Hlginia 
it is hornlike. The anterior part of the sphenethmoid is applied 
to the ventral surfaces of the two frontals. 

Parietal_—The parietals vary in size; sometimes they are larger, 
sometimes smaller than the frontals. The parietal-postparietal 
suture does not generally lie parallel to the frontal-parietal suture, 
but is oblique, with the median end more anterior. On the median 
line in the anterior half of the parietals lies the parietal foramen— 
this is generally rounded but is also sometimes oval; it varies some- 
what in size but is in general small (except in Anthodon), whereas 
in most other Cotylosaurs it is large. Each parietal carries a boss 
with radiating fibres, and in some cases rugae; it lies somewhat 
lateral to the centre of the bone. Ventrally the posterior part of 
the sphenethmoid and the anterior part of the supraoccipital is applied 
to the parietal surface. In one case it would appear that the epiptery- 
goid is also applied to the ventral parietal surface. 

Postparietal—The postparietal differs considerably in size and 
shape, and forms the median portion of the posterior part of the 
dorsal skull-roof. Watson has figured it as paired; only in one case 
have I seen anything that may be a median suture ; in all other cases 
there is, to the contrary, a median boss with radiating bone-fibres 
and sometimes even slight rugae. The posterior edge of this bone is 
of interest in that it may form either a fairly straight, a slightly 
curved, or an emarginate edge, or, again, it may be bent downwards 
to form a flange of bone which in posterior view partly overhangs 
the occiput. This in no way indicates a tendency of the postparietal 
of being incorporated into the occiput itself. On its ventral surface 
it is firmly ankylosed to the ascending strong supraoccipital column. 

Tabular.—The tabular bones are prominent elements in the 
Pareiasaur skull; they form the posterior corners of the dorsal table, 
are provided on their posterior third with a strong boss which in 
some cases is very prominent, and in Elginia forms the longest horn 
on the skull. This boss often has a smaller, more median boss partly 
confluent with it. From the strong tabular bosses well-marked rugae 
run anteriorly, and weaker rugae medially; near the lateral margin 
of the tabular a longitudinal ridge forms the edge of the dorsal table; 
this ridge is sometimes very strong, so that the dorsal and lateral 


10 Annals of the South African Museum. 


surfaces of the skull are sharply separated from each other. The 
posterior edge of the tabular, as in the case of the postparietal, is 
generally a sharp one, but it sometimes helps to form a posterior 
overhanging flange. This flange lies in a plane posterior to the 
occiput; there is therefore no tendency of the dorsal bones entering 
into the formation of the occiput. On its ventral surface abuts the 
strong paroccipital process. The otic notch, at the junction of the 
tabular and squamosal, is practically closed up. 

Lacrymal.—This bone retains its primitive position in that it 
stretches from the orbit to the nostril. It forms an anterior wall to 
the orbit, where it is perforated by a foramen. It is provided with a 
boss, from which the fibres radiate, and in some skulls is strongly 
bulging so that it continues the ridge separating the dorsal from the 
lateral surface of the skull. It does not stretch very far down on the 
lateral surface so as to cause a shallow maxilla to be formed. It 
meets the jugal. In some forms the lacrymal narrows considerably 
as it approaches the nostril. This is not correlated to the geological 
sequence, and it cannot be maintained that there is a tendency towards 
the exclusion of the lacrymal from the nostril. The lacrymal, then, 
retains its primitive Cotylosaurian character. 

Prefrontal.—The prefrontal is a fairly small bone lying on the 
antero-dorsal border of the orbit. It adjoins the nasal, frontal, 
postfrontal, and jugal. Its boss is situated on the dorsal orbital 
border. The prefrontal has a ventrally directed flange, contributing 
partly to the anterior orbital wall, and reaches the upper surface of 
the palatine. 

Postfrontal.—The postfrontal is a fairly small bone lying on the 
dorsal orbital border. In all specimens examined both internal and | 
external sutures show that it is a larger element than figured by 
Watson. In this point Broom is right in his criticism of Watson’s 
figure. It carries a boss on the orbital border, which in some forms 
is confluent with the frontal boss. The suture between this bone 
and the prefrontal lies between the two bosses and joins the frontal 
suture at its middle just where it forms a laterally directed angle. 

Postorbital.—The postorbital is larger than the two foregoing bones. 
It is situated on the dorso-posterior border of the orbit and is pro- 
vided with a strong boss which continues posteriorly as a ridge running 
to the tabular ridge. Together they demarcate the dorsal from the 
lateral surfaces of the skull. This ridge is stronger in some forms 
than in others. It is thus seen that the three supraorbital bosses 
lie each on one of the three supraorbital bones. The lhmiting sutures 


Pareiasaurian Studies.—Part IX. 11 


are generally visible, but, if they are not, one can obtain a very 
accurate idea of the le of the three bones from the three bosses 
with the concomitant radiation of the bone-fibres. | 
Jugal.—The jugal is a large bone forming the ventro-posterior 
border and floor of the orbit. It is provided with a large low boss, or 
a number of low bosses grouped together, from which rugae radiate. 
It sometimes (Bradysaurus vanderbyli, Embrithosaurus schwarzt) 
stretches far posteriorly, thereby encroaching on the quadratojugal. 
In some skulls it is intercalated between the quadratojugal and 
maxilla so that these bones do not meet; in others (Nochelesaurus 
strubeni, Brachypareia watsont) it does not reach the ventral border, 
so that the maxilla just meets the quadratojugal. In most other 
Cotylosaurs (Seymouria, Diadectes, Limnoscelis) the jugal forms a 
large part of the antero-ventral edge of the “cheek,” and the maxilla 
and quadratojugal are thus widely separated. It meets the lacrymal. 
Maxilla.—The maxilla is not such a very shallow bone as Watson 
has maintained. Just back of the nostril it is swollen, and in one 
skull (Pareiasaurus serridens) even has a prominent conical tubercle. 
It is high up on this bulge that the lacrymal-maxilla suture runs 
digitatingly, so that most of the lateral border of the nostril is formed 
by the maxilla. It also forms about half of the ventral floor to the 
nostril. As has been pointed out, it either meets or just does not 
meet the quadratojugal. The outer maxillary surface may be only 
slightly curved so that the teeth are vertical, or (Bradysaurus, Brachy- 
pareia rogerst) 1t may be much curved inwards so that the teeth are 
also directed inwards. In the latter case much of the lateral and 
anterior portions of the palate are obscured in ventral view. In- 
ternally the maxilla sends a thin horizontal plate inwards to form 
part of the bony palate and the anterior part of the lateral border of 
the choanae. The maxilla houses a variable number of teeth. 
Premaxilla.—The premaxilla houses two teeth; forms half of the 
floor to the nostril; sends a process dorsally to form part of the 
internasal bar—this is laterally clasped by a descending nasal process; 
sends a process inwards on to the palate, where it forms the anterior 
border of the choanae; dorsally underneath the internasal bar the two 
premaxillae enclose a foramen—the foramen incisiwum. In Dolicho- 
pareia, although this part is well exposed, no foramen is visible. 
Ventrally, just anterior to the prevomers, three or four small foramina 
have been located—two lie on the median suture (in Anthodon these 
are fused into one), the other two a little lateral to it; all penetrate 
the bone and open on to the dorsal surface. In some cases the 


12 Annals of the South African Museum. 


internasal bar carried a forwardly directed boss (Pareiasuchus nasi- 
cornis). The palatal processes of the premaxilla lie on a higher 
plane than the prevomers, and abut on to the girder formed by the 
anterior ramus of the pterygoid. 

Septomaxilla (fig. 1).—The septomaxilla has been found preserved 
in only one skull (Anthodon). Here it lies wholly within the nostril 
and cannot be seen in dorsal view. It shows no indication of any 
sculpturing as is general in the superficial dermal bones of the skull. 
Its shape is shown in the accompanying figure. 
From the available evidence it appears that 
this bone does not divide the nostril into two 
compartments. 

Squamosal.— There are only two temporal 
bones. Watson has argued in full that these 
two bones can only be the squamosal and 
ine ee ae quadratojugal; Sushkin has claimed to have 

external view. Antho- found another temporal bone in the Russian 

ene ieee !. Pareiasaurs. Hartmann- Weinberg, who has 

subsequently studied this material, could not 

locate this ** supra-temporal.”’ In none of the skulls I have examined 

is there anything that can be considered to form a third bone. I 

have examined the temporal region internally as well as externally 

and have also sections through it, and can definitely state that there 
is no “‘supra-temporal”’ bone present. 

The squamosal is a fair-sized bone with a not-well-defined central 
boss, but with clearly visible radiating bone-fibres and a few rugae. 
The squamosal-tabular suture runs just below the tabular ridge and 
reaches the posterior surface just below the posterior shelf of the 
tabular. Here there is a slight notch—a remnant of the large Stego- 
cephalan otic notch. The posterior border of the squamosal is very 
much thickened and in the majority of forms carries a number of 
bosses, whereas in EHlginia a horn is developed. Internally_ the 
squamosal sends a flange which posteriorly clasps the quadrate; this 
is supplemented by a similar flange from the quadratojugal. The 
squamosal-quadrate, the squamosal-quadratojugal, and the quadrato- 
jugal-quadrate suture meet at a small foramen—the foramen quadratz. 

Quadratojugal.—The quadratojugal forms the angle of the “cheek.” 
At about the centre of its external surface there lies a centre of 
radiation of bone-fibres and in some cases of rugae. This centre is 
in some cases a single boss, but is more often a boss-complex. The 
relative size of the bone differs considerably, and this is also the case 


Se 
Ss ; 


== == 
——— 


Pareiasaurian Studies.—Part IX. 13 


with reference to its shape. It is of interest to note that in those 
forms with high “cheeks” this height is due to increased size of the 
quadratojugal. The angle and posterior border is thickened and is 
sometimes fairly smooth, but more often houses a series of bosses or, 
as an extreme, horns. A study of the nature of the thickening and 
the direction of the bone-fibres seems to indicate that the essential 
structure is the same in the smooth and the bossed forms, and that 
the bosses are simply realisations of potentialities innate in the 
smooth forms. The quadratojugal meets, or just does not meet, the 
maxilla. The quadrate abuts on its ventral surface which posteriorly 
sends a flange, which, in conjunction with a similar flange from the 
squamosal, supports the quadrate from behind. The squamosal- 
quadratojugal and the quadratojugal-quadrate sutures meet at the 
small foramen quadrats. 

Prevomer (figs. 2 and 3).—The prevomer is semicircular in shape, 
the diameter lying medially and is applied to that of its fellow; at 
its diameter it is raised to a ridge carrying a row of small sharp and 
recurved teeth. In some cases (Pareiasuchus) the first two pairs of 
teeth are larger than the rest; along their dorsal surfaces the pre- 
vomers are supported by the median pterygoid girder, which meets 
the palatal processes of the premaxillaries (in some cases the ptery- 
goid girder is very intimately fused to the prevomers, so that, 
even in section, it has not been possible to determine the limits 
of the two bones); medially the anterior borders of the prevomers 
are slightly emarginate. The prevomers differ somewhat in size in 
the different forms, so that the ““comma-shaped”’ vacuities which 
housed the choanae vary in size; in some forms the lateral portions 
of the prevomers do not lie horizontal, so that it appears as if only the 
anterior part of the “‘comma-shaped”’ vacuities could house the choanae. 

Pterygord.—The pterygoids are large elements, together forming 
the greater part of the bony palate, firmly joined to the basisphenoid 
and supporting the quadrate. It is a triradiate bone consisting of 
(a) an anterior flange, (b) a lateral flange, and (c) a posterior flange 
(quadrate ramus). The basisphenoid-pterygoid junction is marked 
by an oblique ridge. From here the anterior flange stretches an- 
teriorly up to the palatal process of the premaxillaries; along the 
median line a girder is developed on the dorsal surface of the ptery- 
goids, and this girder continues anteriorly over the dorsal surface of 
the prevomers to meet the palatal processes of the premaxillaries; pos- 
teriorly this girder does not stretch as far back as the parasphenoidal 
rostrum. (Watson maintained that in his specimen it did.) The 


14 Annals of the South African Museum. 


median edges of the two pterygoids are firmly joined to each other 
along the anterior half of the bone; in the case of two forms (Scuto- 
saurus, Anthodon) the posterior half of the two pterygoids diverge 
posteriorly to leave a fairly long interpterygoid slit (typical of primi- 


Fic. 2.—Dorsal view of the skull of Bradysaurus seeleyx. x 4. S§.A.M., No. 9137, 
with the roof bones removed. The right half of the brain-cavity is exposed. 


A. Pter. =anterior ramus of the | Pr. Vo. = prevomer. 
pterygoid. Pr. Ot. = pro-otic. 
B. Oc. =basioccipital. is Per: = posterior ramus of the ptery- 
B. Sph. =basisphenoid. goid. 
Ch. = choanae. Po Par. = postparietal. 
Ex. Oc. =exoccipital. Qu. = quadrate. 
For. Inc. =foramen incisivum. Qu. Jug. = quadratojugal. 
For. Mag. =foramen magnum. Sub. Orb. nes + 
For. Qu. =foramen quadrati. Vac. ackaiuaieune 
Jug. =jugal. Sub. Temp. : 
L. Pter. Sea ramus of the | Vac. z oe ee 
pterygoid. S. Oc. = supraoccipital. 
Mx. = maxilla. Sq. = squamosal. 
Pal. = palatine. Tab. =tabular. 
Par. Oc. =paroccipital (opisthotic). | Trans. = transversum (ectopterygoid). 
P. Sph. =parasphenoidal rostrum. | T. St. =tubercle for attachment of 
Pr. Mx. =premaxilla. the stapes. 


All illustrations and photographs are, except where otherwise stated, by the 
author. 
tive Stegocephalians and Cotylosaurs) through which the para- 
sphenoidal rostrum is visible; in the other forms the junction of the 
medial pterygoid edges is carried further back so that the slit is 
closed and the interpterygoid space has either a shallow concave, 


Pareiasaurian Studies.—Part IX. 15 


straight, or even convex border through which the parasphenoidal 
rostrum is in some cases partly visible. In these forms where the slit 
is closed it is apparent that the closure has not taken place uniformly, 
viz. in some forms the forward extension of the interpterygoid space 
is greater than in others; one can thus speak of long and short palates. 
The anterior pterygoid flange carries a double row of small, sharp, 


BOc. BSph.  TSt 


Fic. 3.—Ventral view of the skull of Pareiasuchus peringueyt. x +4. S.A.M., 
No. 2337. Drawing by Miss M. van der Merwe. For lettering see fig. 2. 


recurved teeth; these are implanted on the medial pterygoid edge. 
In some forms with a closed interpterygoid slit that has continued 
far posteriorly the teeth do not lie on the medial edge of the part 
thus closed; here a low vault is formed (Bradysaurus, Embrithosaurus). 
In Pareiasuchus peringueyi a short median row of teeth is developed 
on this part of the median line. 

From the basipterygoid junction the lateral flange of the pterygoid 
stretches anteriorly and laterally to meet the palatine and trans- 
versum. Unfortunately these sutures are not often visible. This 
flange of the pterygoid carries (a2) a double row of small, sharp, 
recurved teeth which begins near the basipterygoid junction and 
diverges in antero-lateral direction to where the pterygoid meets the 
palatine, and (6) separated from the foregoing by a deep groove, the 


16 Annals of the South African Museum. 


raised postero-lateral pterygoid edge, which forms the border of the 
suborbital fenestra, sometimes carries a single row of small, sharp, 
recurved teeth (I have not been able to determine this in all skulls, 
and Sushkin also does not figure it in Scutosaurus). In between these 
sets of pterygoid teeth there are deep grooves which carry no teeth; 
the shagreen mentioned by other authors appears to be loose teeth 
from these rows lying scattered in the matrix filling these grooves. 
There is certainly nothing approaching the condition as exemplified 
by Pantylus. 

The posterior pterygoid flange commences at the ridge indicating 
the basipterygoid junction; from here it extends laterally as a more 
or less vertical flange directed somewhat posteriorly. This flange is 
closely applied to the posterior surface of the quadrate; it reaches the 
temporal roof, and was, together with the quadrate, cartilaginously 
attached to the brain-case. Medial to the quadrate-pterygoid suture 
the posterior surface of the bone is more or less deeply excavated. 
In the recess so formed the stapes was probably housed. The 
quadrate sometimes stretches far in median direction along the 
anterior surface of the pterygoid flange, sometimes (Anthodon) 
even meeting the lateral pterygoid flange; in anterior view the 
posterior pterygoid flange was thus wholly or only partly covered 
by the quadrate. It is on the dorsal surface of this ramus of the 
pterygoid, near its junction with the basisphenoid, that the epiptery- 
goid stands. | 

Palatine.—Lateral and anterior to the lateral ramus of the ptery- 
goid lie the palatine and transversum. The palatine, as far as can be 
determined, meets the lateral ramus of the pterygoid where the 
middle double row of teeth ends; from here it curves forward along 
the posterior border of the ““comma-shaped” vacuity and, meeting 
the palatal flanges of the maxilla, forms together with the latter the 
lateral margin of this vacuity. On its dorsal surface, just posterior 
to the ““comma-shaped”’ vacuities, it supports the descending process 
of the prefrontal. 

Transversum.—The transversum lies posterior to the palatine, from 
which it is in part separated by the small lateral palatal foramen 
(foramen palatenum). (A comparison with a similar structure in 
Therocephalians and Gorgonopsians makes it preferable to call this 
foramen the suborbital vacuity.) It forms the lateral part of the 
anterior border of the basitemporal fenestra, the medial part being 
formed by the lateral ramus of the pterygoid. Although the trans- 
versum-pterygoid suture has not been determined in many skulls, 


Pareiasaurian Studies.—Part IX. 17 


it can be stated that it does not stretch far medially of the foramen. 
It is thus quite a small bone and carries no teeth. 

Basioccipital.—The basioccipital forms the whole of the unipartite 
circular articular surface for the vertebral column. Seeley’s erroneous 
account of a tripartite condyle was due to the imperfect nature of the 
specimen examined by him; what he really described was basi- 
occipital plus basisphenoid. In contrast to the condition in most 
Cotylosaurs (except Diadectes and Limnoscelis), the condylar surface 
is concave and is thus similar to the articular surfaces of the vertebrae. 
The bone is massive, short, and is firmly ankylosed to the basisphenoid. 
Through the courtesy of Professor Young of the University of Cape 
Town I have sections through the basioccipital and basisphenoid but 
can find no indication of the extent of these two bones. Watson, 
however, in his figure of the brain-case indicates a suture. 

Basisphenoid.—This is a massive bone with prominent tubera 
on its ventro-posterior angles. Its ventral surface is shallowly or 
deeply grooved; it continues anteriorly as the parasphenoidal 
rostrum, which lies in the interpterygoid space on a plane dorsal to 
that of the palate and meets the sphenethmoid. Laterally the 
basisphenoid is immovably joined to the pterygoid, the line of 
junction being indicated by a sharp ridge. The basisphenoid forms 
the middle portion of the floor of the brain-case; its median upper 
surface is excavated to form the pituitary fossa; laterally it extends 
to meet the pro-otic, and thus forms the median part of the side wall 
of the brain-case; here its outer surface is pierced by the external 
carotid foramen. 

Ezxoccipital.—The exoccipitals form the lateral borders of the 
foramen magnum and a small part of the lateral walls of the posterior 
section of the brain-case. They may or may not, in addition, form the 
upper border of the foramen magnum, in the former case excluding 
the supraoccipital. The dorsomedial corner of the paroccipital process 
is formed by the exoccipital. 

Supraoccipital—The supraoccipital, generally weak in Stegoce- 
phalians and Cotylosaurs, is very strong in the Pareiasaurs; it forms 
a very strong low or high pillar firmly ankylosed to the postparietal; 
it forms the dorsal border of the foramen magnum, or is excluded 
from it; anteriorly to the pillar it sends out a thin flange of bone, 
which forms the side wall of the brain-case right up to the large 
foramen for the Vth nerve; it forms, together with the pro-otic, 
the lower part of the side wall of the brain-case. 

Paroccipital—The paroccipital (or ophisthotic) forms the greater 

VOL. XXXI, PART 1. 2 


18 Annals of the South African Museum. 


part of the paroccipital process, which is essentially a horizontal 
bar of bone connecting the brain-case to the dermal roof-bones. 
It generally abuts on the tabular, but the squamosal may also be 
included; medially it meets the pro-otic, supraoccipital, and ex- 
occipital. The pro-otic forms the antero-medial corner of the par- 
occipital bar, the exoccipital the ventro-postero-medial corner, and 
the supraoccipital the dorso-postero-medial corner. Generally the 
paroccipital forms a strong bar with a slight dorsal curvature; in 
Bradysaurus baint and Pareiasaurus serridens a bulge is developed 
on its dorso-lateral corner. The extent of its curvature, its length, 
and the height of the supraoccipital pillar determine the size and shape 
of the post-temporal fenestra. Its ventral extension also varies so 
that in occipital view the quadrate-pterygoid bar is obscured to a 
greater or lesser extent; 1n some cases it is situated so high up that 
in occipital view the ptero-occipital fenestra is visible. The par- 
occipital forms part of the side wall of the brain-case, viz. that part 
which les between the fenestra ovalis and the foramen jugulare; it 
forms the anterior border of the latter and the posterior border of the 
former. 

Pro-otic.—The pro-otic forms the greater part of the posterior half 
of the side wall of the brain-case; it forms the posterior border of the 
irregular opening for the Vth nerve, the anterior border of the fenestra 
ovalis, the posterior border of the pituitary fossa, and is pierced by a 
small foramen for the VIIth nerve. As already mentioned, it forms 
the antero-medial corner of the paroccipital bar. 

Sphenethmoid (fig. 4).—The whole anterior portion of the brain is 
enclosed by the sphenethmoid, which really merits its French name 
—os en ceinture. It is, however, only its anterior half that wholly — 
encircles the fore-brain; the posterior half, separated from the 
anterior by a ridge on the external surface, is merely a thin flange of 
bone forming the side wall; ventrally it passes into the basisphenoid 
and dorsally it abuts on the frontal, so that the latter—a dermal 
bone—forms that part of the dorsal wall; posteriorly this flange does 
not meet that sent forward by the supraoccipital. Between these 
two flanges the brain has no bony wall; the ventral part of this— 
the posterior sphenethmoidal flange—forms the anterior border of 
the large foramen for the Vth nerve; postero-ventrally the anterior 
part of the sphenethmoid forms the anterior margin of the pituitary 
fossa; between these two openings the sphenethmoid meets the 
ascending process of the pro-otic. A little anterior to and dorsal to 
the pituitary fossa lies a small fissure through which the IInd, [Ird, 


Pareiasaurian Studies.—Part IX. 19 


and IVth nerves emerged; the Ist nerve emerged through the widely 
open end of the sphenethmoid. Ventrally the sphenethmoid is 
supported by the parasphenoidal rostrum. 

From the above account of the bones co-operating to form the brain- 
case, it has become apparent that the Pareiasaur brain-case is long, 


SO POO. 
SK So, 


OA XXX 


Se 


‘i 
iN" 


Fie. 4.—External view of right side of brain-case of Pareiasaurus serridens. x 1. 
The paroccipital process and the posterior pterygoid ramus seen in section 
near their bases. 


Bo. =basioccipital. Pit. =pituitary fossa. 

Bs. =basisphenoid. Po. =paroccipital process seen in section. 
Ee. =external carotid foramen. Pp. =postparietal. 

Ko. =exoccipital. Ppt. =posterior ramus of the pterygoid. 
Ept. =epipterygoid. Pro. = pro-otic. 

Fj. =foramen jugulare. Psr. = parasphenoidal rostrum. 

Fo. =fenestra ovalis. Se. =anterior portion of sphenethmoid. 
Fr. =frontal. Sef. =posterior flange of sphenethmoid. 
Lpt.=lateral ramus of pterygoid. So. =supraoccipital. 

i = parietal. Sof. =anterior flange of the supraoccipital. 
Pf. =pineal foramen. 


Roman numerals refer to the cranial nerves. 


wide, and, in general, low; in some of the younger forms (Parevasaurus) 
it is, however, somewhat higher. The internal ear is situated fairly 
high up on the side wall of the brain-case, and the vestibule has an 
osseous wall separating it in part from the brain-cavity; the fore- 
brain is in primitive fashion enclosed by a sphenethmoid; the relative 
positions of the pituitary fossa and fenestra ovalis appear to vary in 
the different forms. 


20 Annals of the South African Museum. 


Epipterygoid.—The epipterygoid is a fairly thick plate of bone 
standing transversely on the posterior (or quadrate) ramus of the 
pterygoid, near its articulation with the basipterygoid process of the 
basisphenoid; it is not attached to the brain-capsule by any bony con- 
nections, and dorsally it does not extend further than the plane of the 
foramen for the Vth nerve. Ina specimen of Pareiasuchus a remnant 
of bone attached to the ventral surface of the parietal may indicate 
that the epipterygoid did in some forms reach the dermal roof-bones. 

Quadrate.—The quadrate is a large plate of bone standing trans- 
versely in the skull, stretching in dorsal direction from the ventrally 
directed bipartite condyle to the ventral surface of the temporal 
bones, and medially to the posterior or quadrate ramus of the ptery- 
goid; it has no osseous connection with the brain-case. It is firmly 
fixed to the quadratojugal, squamosal, and partly also to the tabular; 
the two former bones send flanges on to its posterior surface, reaching 
inwards to the foramen quadrati, which appears at the junction of 
these sutures, and in addition there appear to be very slight flanges 
on to the anterior surface of the quadrate. The dorso-medial corner 
of the quadrate just touches the antero-lateral corner of the par- 
occipital process; from the medial edge of the quadrate there appears 
to have been, in life, cartilage connecting it to the brain-case. The 
quadrate is, furthermore, immovably connected with the posterior 
ramus of the pterygoid; the transverse quadrate plate stretches 
medially from the condyle, so as to lie anterior to the transverse 
plate of the posterior pterygoid ramus. The medial extent of the 
quadrate varies; in some cases (Anthodon) it actually reaches the 
lateral pterygoid ramus, in others it only reaches half-way of the 
distance between the condyle and the lateral pterygoid flange. In 
anterior view the quadrate thus covers the posterior pterygoid 
ramus wholly or partly. In posterior view the reverse is the case, viz. 
the posterior pterygoid ramus stretches in lateral direction on the 
posterior surface of the quadrate. The pterygoid ramus, which is 
more of a bar than a plate, is deeply excavated on its posterior surface 
to form a recess—the lateral margin of this recess (which probably 
housed the stapes) is also the limit of the pterygoid; the pterygoid- 
quadrate suture runs along this margin to the dorso-medial corner of 
the quadrate plate, where it touches the paroccipital bar; just lateral 
to the pterygoid recess on the medial edge of the quadrate there is: 
situated a tubercle to which the stapes was apparently ligamentously 
attached. The articular condyle of the quadrate is bipartite, the 
inner part being larger than the outer. The quadrate plate, from the 


Pareiasaurian Studies.—Part IX. 21 


temporal bones to the condyle, stands inclined in anterior direction; 
the inclination varies from 95° to 85°. In the primitive condition 
(e.g. Seymouria) the inclination is greater. The quadrate is situated 
further anteriorly in the skull than is the case in most Cotylosaurs 
(except, for instance, Dzadectes). 

Stapes.—The stapes is not preserved in-any skull, but something can 
be determined from the structure of the surrounding bones. Judging 
from the nature of the fenestra ovalis its medial end must have been 
slender. Ifit fitted into the recess in the posterior face of the posterior 
pterygoid ramus it must have been curved and here have been a 
fairly massive bone. For the attachment to the tubercle on the 
quadrate a short process must have been developed, and from here 
it must have curved to form a posteriorly directed process which lay 
in the groove between the end of the paroccipital process and the 
squamosal, and reached the otic notch on the posterior surface 
between the tabular and the squamosal. The stapes was thus 
probably a massive curved bone with two lateral processes and a 
medial process tapering nearly to a point at the fenestra ovalis. 

Teeth.—The teeth (Pl. V) of the upper jaw are carried by the 
maxilla and premaxilla. Two teeth are implanted in the premaxilla, 
whereas a varying number (10-18) is carried by the maxilla. There 
is no differentiation into incisors, canines, and molars, although there 
is some variation between anterior and posterior teeth, viz. the 
former are larger and may have a larger number of cusps. 

A tooth consists of a long root and crown separated, to a greater 
or less degree, by a cingulum. The teeth are thecodont, the roots 
being very firmly held by the encircling bone-tissue. The crown 
is flattened extero-internally (labio-lingually) and elongate antero- 
posteriorly to form, roughly, a semicircular functioning edge. The 
inner surface is flattened or slightly concave; a cingulum separates 
the crown from the root; long and deep grooves radiate from the 
notches between the cusps towards the cingulum. The outer 
surface is convex, imparting a greater or less degree of inflation; 
much shorter and shallower grooves radiate from the notches between 
the cusps. The teeth are more or less crowded, so that in some 
cases the crowns overlap to a greater or less extent, viz. in that the 
posterior edges of the crown overlaps the anterior edge of the suc- 
ceeding tooth. The edge of the crown is developed into a number of 
cusps whose number varies in the different forms, and in some cases 
the posterior teeth have fewer cusps than the anterior ones. In the 
large-bodied forms (Bradysaurus, Nochelesaurus, Dolichopareia, and 


22 Annals of the South African Museum. 


? Bradysuchus) of the Tapinocephalus zone there are 6-8 cusps 
arranged as follows: 1, 2, or 3 anterior, 3 terminal, and 2 or 3 posterior. 

In some of the small-bodied Tapinocephalus-zone forms (Brachy- 
pareia, ? Koalemasaurus) there are also few cusps (6-7) arranged as 
follows: 1 or 2 anterior, 3 terminal, and 2 posterior. 

In Embrithosaurus there is an intermediate number of cusps (9) 
arranged as follows: 3 anterior, 3 terminal, and 3 posterior. 

In the younger forms the cusps become more numerous and are 
more evenly arranged around the crown, so that eventually it becomes 
dificult to maintain the distinction into anterior, terminal, and 
posterior cusps. In Propappus from the Endothiodon zone there are 
10 cusps—3 anterior, 4 terminal, and 3 posterior. In Pareiasuchus 
(Cistecephalus zone) there are 11-13 cusps fairly evenly arranged. 
In Parevasaurus there are 11 fairly evenly arranged cusps. In 
¢ Pareiasaurus russouwi there are up to 15 cusps evenly arranged. 
In Anthodon there are up to 15 cusps evenly arranged. In the 
Russian Scutosaurus the number of cusps according to Hartmann- 
Weinberg vary up to a maximum of 17, and presumably they are 
evenly arranged. In the Scottish Hlginia there are, according to 
Newton, 11 cusps evenly arranged. 


MEASUREMENTS. 


All but one of these measurements have been taken by Martin’s 
beam-compass and are therefore direct length, 2.e. projections on to 
a horizontal or vertical plane. Only the dentigerous length has been 
taken by means of a tape-measure. Where more than one specimen 
of a species have been measured, the figures given are the mean. _ 

1. Median length, premaxillary suture to basioccipital condyle. 

2. Length from anterior border of pineal foramen to premaxillary 
suture. 

3. Length from posterior border of pineal foramen to posterior 
margin of postparietal. 

4. Width, across angles of “cheeks.” 

5. Intertabular width, distance between tops of tabular bosses. 

6. Interorbital width, minimum distance between dorsal orbital 
borders. 

7. Length from anterior orbital border to premaxillary suture. 

8. Length of “cheek,” minimum distance from posterior border 
of orbit to posterior border of cheek. 

9. Height of “cheek,” top of tabular boss to angle of “cheek.” 


23 


“SOLPOUTT[IUL UL VIB SJUSWIOINSVOU OSO44 [[V 


Pareiasaurian Studies.—Part IX. 


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24 Annals of the South African Museum. 


10. Overhang of “‘cheek,”’ distance of vertical projection of “‘cheek”’ 
below the horizontal level of dentigerous maxillary border. 

11. Palatal width, distance across last teeth. 

12. Palatal length, anterior border of prevomer to anterior border 
of interpterygoid vacuity. 

13. Interquadrate width, distance between lateral edges of the 
condyles. 

14. Dentigerous length, from premaxillary suture to last tooth. 

15. Number of teeth. 

16. Angle of inclination of the quadrate to the plane of the palate. 

17. Postparietal: (a) length; (6) width. 


Genus BRADYSAURUS Watson. 


This genus includes forms with large skulls; forms with many 
(19-20), and forms with an intermediate number (15-16) of teeth; 
these teeth may have no, slight, or considerable overlap. All the 
forms have teeth with few cusps (less than 9) arranged as follows: 
2 anterior, 3 median, and 1 or 2 posterior; the teeth are moderately, 
or very much, swollen. The interorbital width is appreciably less 
than the “tabular width.”” The posterior border of the “cheek” is 
very smooth, fairly smooth, or has well-defined bosses. The snout is 
broad and rounded, and the “cheek” shallow. In general the skull 
is fairly smooth or is ornamented with pits and deep rugae. The 
maxilla and lacrymal are only slightly, or very strongly, bulging. 
Both behind and in front of the orbit the dorsal and lateral surfaces 
are separated by a strong ridge, or by only a moderate ridge. The 
postparietal is small to large, square or longitudinally or transversely 
rectangular. Medially the pterygoids are fused far posteriorly to 
form a long palate; the border of the interpterygoid vacuity is either 
concave, straight, or convex; and anterior to the border, in between 
the median rows of pterygoid teeth, there is a shallow vault. The 
quadrate is inclined forwards (70°-85°) and does not stretch far 
medially, so that in anterior view the quadrate ramus of the pterygoid 
is partly exposed. The brain-case is fairly low; so is the supra- 
occipital pillar; the supraoccipital is excluded from the foramen 
magnum. The tabular boss is either low or very prominent. The 
maxilla is fairly shallow, and the quadratojugal shallow or fairly 
deep. The jugal just separates the maxilla from the quadratojugal. 

In their preliminary classification Haughton and Boonstra have 
placed a distorted skull (S.A.M., No.5002) in a new genus—Platyoropha. 
Subsequent study of the postcranial skeleton has proved that this 


Pareiasaurian Studies.—Part IX. 25 


specimen does not differ much from the forms referred to the genus 
Bradysaurus. A re-examination of the skull has, moreover, shown 
that the features considered different are in all probability due to 
the post-mortem deformation that the skull has suffered. Making 
allowance for this, and taking into consideration that there are no 
other characters, either in the skull or in the postcranial skeleton, 
which could exclude this specimen from the genus Bradysaurus, itis best 
referred to that genus. It most closely approaches the species bani. 
Platyoropha broom: thus becomes a synonym of Bradysaurus baini. 

Bradysaurus bombidens was founded by Owen on a very fragmentary 
maxilla and mandible. This very unsatisfactory type was founded 
mainly on the nature of the teeth. The inflation and inward direction 
of the teeth is, however, also found in other forms. The specimen 
is therefore specifically indeterminate, but is evidently very close to 
the large-bodied forms of the Tapinocephalus zone. 


Bradysaurus baini (Seeley). 
(Pls. I-V.) 


In this species there are an intermediate number of teeth (15-16), 
which slightly overlap, have 7 cusps, and are only slightly swollen. 
The posterior border of the “cheek” is fairly smooth, but the general 
surface of the skull is strongly rugose and pitted. The maxilla and 
lacrymal are very strongly bulging. Both behind and in front of 
the orbit the dorsal and lateral surfaces of the skull are sharply 
separated by a strong ridge. The postparietal is large and trans- 
versely rectangular. The anterior border of the interpterygoid is 
convex. The tabular boss is low. 

The following skulls are assigned to this species :— 


Type, Brit. Mus. Nat. Hist., No. R1971, Tapinocephalus zone, high. 


South African Museum, No. 3533 & See later: 
ue a 4347 a a) low: 
i . 4351 0 Lowe: 
Y i 4999 " high. 
i a 5000 fe low. 
- ah 5002 we a ne: 
F a 5127 a Stra 
a a 9001 as ad 
is x3 9104 r elon; 
9105 Be ee lows 


Geol. Pal. Inst. Tiibingen, 4 lows 


26 Annals of the South African Museum. 


Bradysaurus seeley: (Haughton and Boonstra). 
(Pls. I-V and text-fig. 2.) 


In this species there are many teeth (19-20), which overlap con- 
siderably, have 7 cusps, and are much swollen. The posterior border 
of the ““cheek”’ has strong, well-defined bosses. In general the skull 
is rugose and somewhat pitted. The maxilla and lacrymal are 
bulging. The dorsal and lateral surfaces are separated by a low 
ridge weaker than in baini. The postparietal is of moderate size and 
is roughly square. The anterior border of the interpterygoid vacuity 
is nearly straight. The tabular boss is very prominent. 

The following skulls are assigned to this species :— 

Type, British Museum, No. 49426 (Palmiet Specimen), Tapino- 

cephalus zone, high. 

Co-type, British Museum, No. R1970 (Tamboer Specimen), Tapino- 

cephalus zone, mid. 
South African Museum, No. 1204, Tapinocephalus zone, mid. 


5 ~ 4346 ie . 1oWw 
o 5624 a » low. 
ML s 8933 ss +5 
‘. ¥; 9115 _ 3 mmid: 
- 3 9127 M 550 KORN 
* 9121 = >»  anad 
.: m 9137 a 3. med? 
* “sf 9168 re «5. mde 
Durban Museum, Pa » low. 


Bradysaurus vanderbylt (Haughton and Boonstra). 
(Pls. I-V.) 


There are an intermediate number of teeth (15-16), which do not 
overlap, have 7 cusps, and are moderately swollen. The posterior 
border of the “cheek” is very smooth, and so is the general surface 
of the skull. The maxilla and lacrymal are slightly bulging. The 
dorsal and lateral surfaces are not sharply separated as the ridge is 
low and rounded. The postparietal is small and is longitudinally 
rectangular. The anterior border of the interpterygoid vacuity is 
concave. The tabular boss is prominent. 

The following skulls are assigned to this species :— 

Type, South African Museum, No. 3718, Tapinocephalus zone, low. 

6 . 8941 _ 35. pene 
i " 9169 5 sof, lands 


Pareiasaurian Studies.—Part IX. 7a || 


Genus BrapysucHus Haughton and Boonstra. 
Bradysuchus whartst (Broom). 
(Pls. I-II.) 


As I have not had the opportunity of seeing the type specimen in 
the American Museum of Natural History, this diagnosis had to be 
based on photographs kindly supplied by that institution and also 
on information received from Dr. R. Broom. 

The skull of the single specimen of this form is large; the teeth are 
badly preserved, but there were probably about 16 teeth with few 
(less than 9) cusps, probably arranged as in the genus Bradysaurus; 
the interorbital width is appreciably less than the “tabular width.” 
The posterior border of the “‘cheek”’ carries well-defined bosses. In 
general the skull is ornamented with pits and rugae. The maxilla 
and lacrymal are only slightly bulging. Both behind and in front of 
the orbit the dorsal and lateral surfaces are separated by a strong 
ridge. Medially the pterygoids are fused far posteriorly to form a long 
palate. The quadrate is inclined forwards. The brain-case and the 
supraoccipital pillar appear to be high. The tabular boss is 
prominent. 

Except for Broom’s statement that the lower jaw carries two 
bosses on the angular, and the fact that the snout appears to be 
somewhat pointed, this genus is very similar indeed to the forms 
included in the genus Bradysaurus, particularly to Br. seeleyt. 


Type, Amer. Mus. Nat. Hist., No. 5567, Tapinocephalus zone, low. 


Genus NocHELESAURUS Haughton and Boonstra. 


This genus includes forms with fairly large skulls; forms with an 
intermediate (15-17) number of teeth—these teeth may have a slight 
or a considerable overlap; all forms have teeth with few cusps (less 
than 9) arranged as follows: 3 anterior, 3 median, and 2 posterior; 
they are slightly swollen. The interorbital width is approximately 
equal to the “tabular width.” The posterior border of the “cheek”’ 
is smooth, or carries very strong bosses. The snout is rather pointed 
and the “cheek” deep; the latter is vertical or somewhat diverging. 
The skull is in general very smooth or strongly ornamented, but 
with little “crocodilian” pitting. The maxilla and lacrymal are 
moderately or slightly bulging. The dorsal and lateral surfaces are 
not separated by a very strong ridge. The postparietal is either 
transversely or longitudinally rectangular. The pterygoids are fused 


28 Annals of the South African Museum. 


far posteriorly so as to form a long palate. The border of the inter- 
pterygoid vacuity is concave. There is hardly any vault between 
the medial rows of pterygoid teeth. The quadrate is much (55°), or 
slightly (85°), inclined forwards and does not stretch far medially. 
The brain-case is fairly low; so is the supraoccipital pillar. The 
tabular boss is low. The maxilla is fairly deep and the quadratojugal 
moderately deep. The quadratojugal just meets the maxilla or is 
separated from it by the jugal. 


Nochelesaurus strubeni (Broom). 
(Pls. I-V.) 


The teeth are slightly overlapping; the whole skull is smooth; 
there is a single nasal boss; the posterior border of the “cheek” has 
hardly any development of bosses. The snout is higher and more 
pointed than in N. alexanderi, and the “cheeks” are more vertical. 
The maxilla and lacrymal are slightly bulging. The postparietal is 
longitudinally rectangular. The quadrate is slightly inclined forwards 
(85°). The quadratojugal just meets the maxilla. From his descrip- 
tion and figures it appears that von Huene is right in assigning the 
lower jaw in the Tiibingen collection to this species. 

The following skulls are assigned to this species:— 

Type (lower jaw): 

Amer. Mus. Nat. Hist., Tapinocephalus zone, low. 

South African Museum, No. 1207, Tapinocephalus zone, low. 
e 4352 5 9 Ws 
2: - 5019 3 aie 
B 2 5590 53 es 

Geol. Pal. Inst. Tibingen, a 3 ee 


Nochelesaurus alexanderi Haughton and Boonstra. 
(Pls. I-V.) 


The teeth are considerably overlapping; the whole skull is heavily 
ornamented with bosses and rugae; there are two nasal bosses; 
there is a great development of bosses on the posterior border of the 
“cheek” approaching the condition shown in Pareiasuchus perin- 
gueyt. The snout is lower and broader than in WN. strubeni, and the 
“cheek” is more flattened out. The maxilla and lacrymal are 
moderately bulging. The postparietal is transversely rectangular. 


Pareiasaurian Studies.—Part IX. 29 


- ‘The quadrate is much inclined forwards (55°). The jugal is inter- 
calated between the quadratojugal and the maxilla. 
The following skulls are assigned to this species:— 


Type, South African Museum, No. 6239, Tapinocephalus zone, mid. 
6239A s r 
39 3) 8944 99 3) 99 


Genus EMBRITHOSAURUS Watson. 
Embrithosaurus schwarz Watson. 
(Pls. I-V.) 


In this genus the skull is quite large; there are an intermediate 
number of teeth (15-16), which are considerably overlapping; there 
are an intermediate number of cusps (9) arranged as follows: 3 
anterior, 3 median, and 3 posterior; they are slightly swollen. The 
interorbital width is approximately equal to the “tabular width.” 
The posterior border of the “cheek” carries pronounced bosses. 
The snout is rather pointed and the “cheek” deep. In general the 
skull is ornamented with pits and rugae. The maxilla and lacrymal 
are only very slightly bulging. The postparietal is transversely 
rectangular. Medially the pterygoids fuse far posteriorly to form a 
long palate; the border of the interpterygoid vacuity is convex. 
The quadrate has a moderate forward inclination (75°), and it does 
not stretch far medially. The brain-case and supraoccipital pillar 
are fairly low. The tabular boss is prominent. The maxilla is 
fairly deep, but the quadratojugal is shallow and small. The jugal 
is intercalated between the quadratojugal and the maxilla. The 
mandible and teeth with 9 cusps described and figured by Broom must 
be assigned to this species if we are correct in maintaining that the 
nature of the cusping is a feature of taxonomic importance. 

The following skulls are assigned to this species :— 


Type, South African Museum, No. 8034, Tapinocephalus zone, mid. 
Brit. Mus. Nat. Hist., 3 5 dae, 
(Watson’s Hottentots River Specimen.) 


Genus DoLticHoPaREIA Haughton and Boonstra. 
Dolichopareia angusta Haughton and Boonstra. 
(Pls. I-V.) 


In this genus the skull is long and fairly narrow; there are many 
teeth (19-20), which are slightly overlapping; there are few cusps 


30 Annals of the South African Museum. 


(less than 9) arranged as follows: 3 anterior, 3 median, and 2 pos- 
terior; they are slightly swollen and decrease much in size in antero- 
posterior direction. The interorbital width is approximately equal 
to the “tabular width.’ The posterior border of the ‘“‘cheek”’ carries 
strong bosses. The snout is very pointed and the “cheek” deep. 
In general the skull is moderately ornamented with pits and slight 
rugae. The maxilla and lacrymal are only slightly bulging. Pos- 
terior to the orbit a strong ridge separates the dorsal from the ventral 
surface of the skull. The postparietal is irregularly shaped and its 
posterior border is notched. Medially the pterygoids are fused fairly 
far posteriorly so as to form a moderately long palate. The border 
of the interpterygoid vacuity is convex. The quadrate is much 
inclined (60°) and is not directed medially but anteriorly (see Pl. III, 
fig. 14). The brain-case and supraoccipital pillar are fairly high, the 
supraoccipital is not excluded from the foramen magnum. The 
tabular boss is fairly prominent. The maxilla is fairly deep and so 
is the quadratojugal. The quadratojugal just fails to meet the 
maxilla. 
The following skulls are assigned to this species:— 


Type, South African Museum, No. 6238, Tapinocephalus zone, mid. 
3717 9 ee: 


33 39 


Genus KoALEMASAURUS Haughton and Boonstra. 
Koalemasaurus acutirostris (Broom). 
(Pl. III and text-fig. 5.) 


Through the kindness of Mr. J. Hewitt, of the Albany Museum, I 
have been able to study Broom’s type, which consists of a little more 
than the half of the mandible with a very imperfect skull (parts of 
the cheek, upper jaw, palate, and basis cranu alone are preserved 
in a bad condition). As is the case in so many of Broom’s types, 
this type is really very bad, and it is with reluctance that an attempt 
is made to place it in this system of classification. In our preliminary 
classification we accepted Broom’s identification of the specimen in 
the American Museum of Natural History and based our diagnosis 
on both specimens. Through the courtesy of the American Museum 
I have photographs of the specimen in that institution and am able 
to confirm Broom’s identification. Although both skulls are not 
very well preserved the position of the orbit and the nature of the 
angular boss justifies such an identification. The skull in this genus 
is of moderate size; there are an intermediate number (15-16) of 


Pareivasaurian Studies.—Part IX. Bil 


teeth with a fair amount of overlap; the teeth have few cusps (less 
than 9) arranged as follows: 2 anterior, 3 median, and 1 or 2 pos- 
terior. The interorbital width is greater than the “tabular width.” 
The posterior border of the “cheek” carries low but distinct bosses. 
The snout is high and pointed (but in the American Museum specimen 
it appears to be truncated). In general the surface of the skull is 
moderately ornamented. The maxilla and lacrymal are moderately 
bulging. The dorsal and lateral surfaces are only distinctly demar- 
cated by a ridge behind the orbit. The postparietal is transversely 
rectangular. The pterygoids are fused far anteriorly so as to form a 
short palate; the border of the interpterygoid vacuity is convex. 
The quadrate is inclined forwards (75°) and is situated far forward in 
the skull. The brain-case and supraoccipital pillar are fairly high. 
The tabular boss is fairly prominent. The maxilla is fairly shallow 


Fic. 5.—External view of lower jaw of Koalemasaurus acutirostris. x 4. 
Type in Albany Museum. 


Ang. =angular. Dent. =dentary. 
Art. =articular. Sp. =splenial. 
Cor. =coronoid. Sur Ang. =surangular. 


and the quadratojugal deep. The orbit is in the anterior half of the 
skull. As the angular boss is such a characteristic feature a figure of 
the lower jaw is included (fig. 5). The following skulls are assigned 
to this species :— 


Type, Albany Museum, Tapinocephalus zone, high. 
American Museum, No. 5568, ks a 2 
South African Museum, No. 4345, Br op  Litiealay 


Genus BRacHYPAREIA Haughton and Boonstra. 


Both species of this genus have medium-sized skulls; an inter- 
mediate (15-16) number of teeth, which are slightly overlapping; 
there are few cusps (less than 9) arranged as follows: 3 anterior, 
3 median, and 2 posterior—they are slightly swollen. The inter- 
orbital width is appreciably greater than the “tabular width.” The 
snout is either rounded or rather pointed; the “cheek” is broad and 


32 Annals of the South African Museum. 


low, and its posterior border is rather smooth or has fairly marked 
bosses. The orbit does not lie in the anterior half of the skull. In 
general the skull is fairly smooth, with a slight indication of 
pitting and rugae. The maxilla and lacrymal are slightly bulging, 
more so in rogerst than in watsont. Medially the pterygoids fuse far 
anteriorly, thus producing a short palate; in one species the overhang 
of the maxillaries and premaxillaries increase this appearance of 
shortness. The quadrate is not inclined much forwards (80°) and 
it does not stretch far medially. The brain-case and supraoccipital 
pillar are low. The tabular boss is low. The maxilla and quad- 
ratojugal are fairly deep. The jugal is intercalated between the 
quadratojugal maxilla. 


Brachypareva rogers: (Broom). 
(Pls. I-III, V.) 


The snout is rather rounded and depressed; the posterior border 
of the “cheek” is rather smooth. The maxillaries and premaxillaries 
overhang the anterior part of the palate, so that in ventral view the 
“comma-shaped” vacuities are partly obscured. 

The following skulls are referred to this species :— 


Neotype, South African Museum, No. 5012, Tapinocephalus zone, low. 


9 +B) 4350 e) 9 mid. 
9 oy) 9095 3 9 low. 
? 9 9098 ” ” mid. 


Brachypareia watsont Haughton and Boonstra. 
(Pls. I-II, IV—V.) 


The snout is rather pointed and high; the surface of the “cheek” 
is rugose and carries fairly prominent bosses on its posterior border. 
There is hardly any overhang of the maxillaries and premaxillaries. 


Type, South African Museum, No. 6240, Tapinocephalus zone, low. 


Genus Propappus Seeley. 
Propappus omocratus (Seeley). 
Propappus parvus (Haughton). 

(PIV) 


In Propappus omocratus only a fragment of maxilla and premaxilla 
is known. The lower jaw has two angular bosses; there are an 


Pareiasaurian Studies.—Part IX. ao 


intermediate number of teeth (14), which do not overlap; there 
are many cusps (10-11) arranged evenly in a semicircle. Judging 
from the rest of the skeleton, the skulls must be of medium size in 
omocratus and rather small in parvus. 


Type, South African Museum, No. 1058, Endothiodon zone. 


Genus ParErasucHus Broom and Haughton. 


In this genus the skull is of medium size; there are few teeth (14), 
which do not overlap; there are many cusps (more than 9) arranged 
evenly around the edge of the crown. The interorbital width is 
approximately equal to the “tabular width”; the snout is very broad, 
depressed, and rounded; the “cheek” is deep and carries strong or 
very strong bosses on its posterior border. In general the skull is 
rugose and pitted, more so in peringueyi than in nasicornis. The 
maxilla and lacrymal are bulging. The postparietal is small and 
transversely rectangular. The dorsal and lateral skull surfaces are 
not separated by aridge. Medially the pterygoids fuse far forward so 
as to produce a short palate. The quadrate is inclined much forwards 
(55°-65°) and does not stretch far medially. The brain-case and 
supraoccipital pillar are low; the opening of the Vth nerve is low. 
The supraoccipital forms the dorsal border of the foramen magnum. 
The tabular boss is low. The maxilla is shallow, but the quadrato- 
jugal is large and deep; it does not reach the maxilla. 


Pareiasuchus peringueyt Broom and Haughton. 
(Pls. I-V.) 


There are very strong bosses on the posterior border of the “cheek,”’ 
whose surface is also rugose. The quadrate is very much inclined 
forwards (55°). The angle of the “cheek” is situated far forward. 
The nostrils are visible in dorsal view. 

The following skulls are assigned to this species :— 

Type, South African Museum, No. 2337, Cistecephalus zone, low. 


- ¥ 2367 ‘ 
x s 10668 


a LOM 
29 29 mid. 


Pareiasuchus nasicorms Haughton and Boonstra. 
(Pls. I-III.) 


There are strong bosses on the posterior border of the “cheek,” 


whose surface is fairly rugose. The quadrate is inclined forwards 
VO. .xXxt, PART i. a 


34 Annals of the South African Museum. 


(65°). The angle of the “cheek” is situated far back. Owing to 
the forward direction of the nasal bosses the nostrils are not visible 
in dorsal view. 


Type, South African Museum, No. 3016, Endothiodon zone. 


Genus PAREIASAURUS Owen. 


Pareiasaurus serridens Owen. 


(Pls. I-V.) 


The skull is medium sized; there are few teeth (14), which overlap 
considerably; the number of cusps varies—9 on the anterior and up 
to 11 on the posterior teeth; these cusps are regularly arranged 
around the crown and are of equal size. The interorbital width is 
less than the “tabular width.” The “cheek” is very deep and 
carries strong bosses. The snout is high and pointed. In general 
the skull is fairly rugose and is considerably pitted. The maxilla 
and lacrymal are only slightly bulging. The dorsal and lateral skull 
surfaces are not separated by a ridge but pass into each other in 
rounded curves. The tabular and postparietal have flange-like 
developments overhanging the occiput. Medially the pterygoids 
are fused far anteriorly so as to form a short palate. The border of 
the interpterygoid vacuity is concave, and the parasphenoidal rostrum 
is partly visible through it. The quadrate is nearly vertical, the 
forward inclination being 85°; it also stretches far medially so that 
in anterior view it nearly obscures the posterior pterygoid ramus 
altogether. The brain-case and the supraoccipital pillar are high, 


the fenestra ovalis is small, and the opening for the Vth nerve is 


situated high up on the side wall. The supraoccipital forms the 
dorsal border of the foramen magnum. The maxilla and quadrato- 
jugal are deep, and the jugal is intercalated between the maxilla and 
quadratojugal. 

The following skulls are assigned to this species :— 

Type (cast), Brit. Mus. Nat. Hist., No. R4063, Cistecephalus zone, 

mid. 

Neotype, University of Stellenbosch, Cistecephalus zone, high. 

As only a part of the lower jaw and some teeth are known of the 
specimen, which Seeley named Parezasaurus russouwi (Brit. Mus., No. 
R1996), it is not possible to include it in our scheme with certainty. 
The nature of the teeth is, however, so similar to that of Parevasaurus 
serridens that there appears to be little doubt as to the generic 


Pareiasaurian Studies.—Part IX. 35 


name, but it appears advisable to retain Seeley’s specific name— 
thus Pareiasaurus russouwi. Recently I collected another mandible 
(S.A.M., No. 10667) fairly high up in the Cistecephalus zone which 
I refer to the genus Pareiasaurus. 


Genus ANTHODON, Owen. 


Anthodon serrarvus, Owen. 


(Pls. I-III, V.) 


The skull is small; there are few teeth (11-14), which do not 
overlap; there are many cusps (8-15) of equal size arranged regularly 
around the crown. The interorbital width is less than the “tabular 
width.” The posterior border of the “cheek” has small and low 
bosses. The snout is fairly high and rather pointed; the “cheek”’ 
is quite deep. In general the skull is moderately ornamented with 
pits and rugae. The maxilla and lacrymal are only very slightly 
bulging, but there is a small prominent tubercle on the maxilla. 
The dorsal and lateral surfaces are not sharply separated by a ridge. 
Medially only the anterior half of the pterygoids fuse, so that pos- 
teriorly a long interpterygoid slit remains open. The quadrate is 
moderately inclined forwards (75°), but it extends far medially to 
meet the lateral pterygoid flange so that in anterior view the posterior 
pterygoid ramus is altogether covered by the quadrate. The brain- 
case and supraoccipital pillar are high. The tabular boss is low. 
The maxilla and quadratojugal are deep. The jugal separates the 
maxilla from the quadratojugal. The pineal foramen is very large. 

The following skulls are assigned to this species :— 


Type, Brit. Mus. Nat. Hist., No. 47337, Cistecephalus zone, mid. 


Amer. Mus. Nat. Hist., No. 7001 as 54) Taal 
South African Museum, No. 4020 ms ss Lowe 
45 a 7841 i a. low: 
10074 a lowe 


Owen’s type of serrarius is an imperfect skull from which Watson 
has, however, been able to determine a number of features. I have 
described a lower jaw of a specimen in the South African Museum 
(S.A.M., No. 4020) as of this species. This specimen has also a 
partial skull of which I have been able to determine the whole palatal 
structure. At the end of 1930 Broom published two photographs and 
a short description of a specimen in the American Museum, which 
he maintains is certainly a distinct species—gregoryi. In April 1931 


36 Annals of the South African Museum. 


I found an impartial skull and some elements of the postcranial 
skeleton of an Anthodon (S.A.M., No. 10074) at Dunedin. The 
pelvis and part of the hind-limb have been described without a 
specific determination, as I then doubted the validity of Broom’s 
new species. Measurements of these four specimens show a uniform 
progression from a small form (Owen’s), through two slightly larger 
forms (Broom’s and 8.A.M., No. 4020), to another slightly larger form 
(S.A.M., No. 10074). But they do not differ in any essentials of struc- 
ture, and in Broom’s description of his new species he gives no reasons 
for believing his specimen to differ from Owen’s. He simply states 
that it certainly is a distinct species. In view of these considerations 
it is best to refer them all to Owen’s Anthodon serrarius, and refer the 
slight variation in size to a variation within the species or, maybe, 
an age variation. A complete mandibular ramus in the Geological 
Museum of the University of Stellenbosch is also assigned to this 
species. Anthodon minusculus, from Tanganyika, is only known 
from the distal ends of two humeri. 


Genus ScurosauRuS Hartmann- Weinberg. 


Scutosaurus karpinskw Amalitzky. 
(Pls. I, III-IV.) 


The skull is very large; there are a variable number of teeth with 
many cusps (9-17) arranged regularly around the crown. The 
interorbital width is much less than the “tabular width.” The 
posterior border of the “cheek” carries well-defined bosses. The 
snout is fairly broad and deep, and the “cheek” is deep. In general 
the skull is sculptured with pits and rugae. The maxilla and lacrymal 
are bulging. The dorsal and lateral surfaces are not abruptly demar- 
cated by a ridge but pass into each other in rounded curves. Medially 
the pterygoids are only fused along their anterior half, so that pos- 
teriorly a long interpterygoid slit remains through which the para- 
sphenoidal rostrum is visible. According to Hartmann-Weinberg the 
quadrate is nearly vertical, but according to Sushkin inclines at 
45°, and it does not stretch far in medial direction. The brain-case 
and supraoccipital pillar are high. The tabular boss is low. 


Type, mounted specimen, Geol. Mus. Acad. Sci., Leningrad. 


In Amalitzky’s memoir “diagnoses” are given of three further 
species, but as the figures are so poor, the text so meagre, and other 
Russian authors do not mention them, it would be foolhardy to 


Pareiasaurian Studies.—Part IX. BT 


attempt to establish their systematic position. It would, however, 
appear that one is justified in referring them all to the genus Scuto- 
saurus, for they manifestly cannot be included in the South African 
genus Pareiasaurus. 


Genus Exainta Newton. 


—Elginia mirabilis Newton. 
(Pls. J-II.) 


The skull is small; there are few teeth (12), which do not overlap: 
there are many cusps (9-10) of equal size arranged regularly around 
the crown. The interorbital width is less than the “tabular width.”’ 
The posterior border carries horns. The snout is high and fairly 
pointed, and the “cheek” is deep. In general the skull is greatly 
ornamented, carrying a great development of horns, in some respects 
of an analogous nature to that in the living Moloch and Phrynosoma. 
The maxilla and lacrymal carry protruding horns. The quadrate 
appears to be much anteriorly inclined (45°). The maxilla is deep. 
A long and strong horn is developed on the tabular, and represents 
the tabular boss. 


Type, Coll. Geol. Survey, Elgin Sandstone. 


LITERATURE CITED. 


AMALITZKY, V.—‘‘ Diagnoses of the New Forms of Vertebrates and Plants from 
the Upper Permian on North Dvina,” Bull. Acad. Sci. Russ., 1922. 

Boonstra, L. D.—‘‘Pareiasaurian Studies. Part III.—On the Pareiasaurian 
Manus,” Ann. 8. Afr. Mus., vol. xxviii, pt. i, 1929. 

Boonstra, L. D.—‘‘Pareiasaurian Studies. Part IV.—On the Pareiasaurian 
Pes,” Ann. S. Afr. Mus., vol. xxviii, pt. i, 1929. 

Boonstra, L. D.—‘‘A Contribution to the Cranial Osteology of Pareiasaurus 
serridens Owen,” Ann. Univ. Stellenbosch, vol. viii, Section A, No. 5, 1930. 
Boonstra, L. D.—‘‘Pareiasaurian Studies. Part VII.—On the Hind Limb of 
Two Little-known Pareiasaurian Genera: Anthodon and Pareiasaurus,” Ann. 

S. Afr. Mus., vol. xxviii, pt. iv, 1932. 

Boonstra, L. D.—‘‘Pareiasaurian Studies. Part VIII.—The Osteology and 
Myology of the Locomotor Apparatus.—B. Fore Limb,” Ann. 8S. Afr. Mus., 
vol. xxviii, pt. iv, 1932. 

Boonstra, L. D.—‘‘A Note on the Hyoid Apparatus of Two Permian Reptiles 
(Pareiasaurians),’ Anat. Anz., Bd. 74, 1932. 

Boonstra, L. D.—“ A Study in Evolution: The Phylogenesis of the Parevasauridae,”’ 
S. Afr. Journ. Sci., vol. xxix, 1932. 

Boonstra, L. D.—‘‘Paleobiologiese Beskouinge oor ’n Uitgestorwe Reptielgroep 
(Pareiasauridae),” S. Afr. Journ. Sci., vol. xxix, 1932. 


38 Annals of the South African Museum. 


Broom, R.—‘On an Almost Perfect Skeleton of Pareiasaurus serridens Owen,” 
Ann. 8S. Afr. Mus., vol. iv, pt. ii, 1903. 

Broom, R.—‘‘ A Comparison of the Permian Reptiles of North America with those 
of South Africa,” Bull. Amer. Mus. Nat. Hist., vol. xxviii, pp. 197-234, 1910. 

Broom, R.—“On Four New Fossil Reptiles from the Beaufort Series, South 
Africa,’ Rec. Alb. Mus., vol. ii, 1913. 

Broom, R.—‘“‘ Further Observations on the South African Fossil Reptiles,’ Amer. 
Mus. Journ., vol. xiv, 1914. 

Broom, R.—‘‘Catalogue of Types and Figured Specimens of Fossil Vertebrates in 
the American Museum of Natural History. JJ.—Permian, Triassic, and 
Jurassic Reptiles of South Africa,’ Bull. Amer. Mus. Nat. Hist., vol. xxv, 1915. 


Broom, R.—“ Pareiasaurian Nomenclature,” Ann. Mag. Nat. Hist., ser. viii, vol. 
xvii, 1916. 
Broom, R.—‘On Some Points in the Structure of the Pareiasaurian Skull,” 


P.Z.S., vol. i, 1924. 
Broom, R.—‘‘ On a New Species of Anthodon (A. gregoryi),” Amer. Mus. Novitates, 
No. 448, 1930. 


Broom, R., and Haucuton, 8. H.—‘‘On the Skeleton of a New Pareiasaurian 
(Pareiasuchus peringueyt),’’ Ann. 8. Afr. Mus., vol. xii, pt. i, 1913. 

HarRTMANN-WEINBERG, A.—“‘Ziir Systematik der Nord Diina Pareiasauridae,”’ 
Palaeontologische Zeitschrift, Berlin, Bd. 12, 1930. 

Havucuton, S. H.—‘‘Pareiasaurian Studies. Part I].—Notes on Some Pareia- 
saurian Brain-cases,’’ Ann. 8. Afr. Mus., vol. xxviii, pt. i, 1929. 

Havucuton, S. H., and Boonstra, L. D.—‘‘ Pareiasaurian Studies. Part I.—An 


Attempt at a Classification of the Pareiasauria based on Skull Features,” 
Ann. 8. Afr. Mus., vol. xxviii, pt. i, 1929. 

HavueutTon, S. H., and Boonstra, L. D.—‘‘ Pareiasaurian Studies. Part V.—On 
the Pareiasaurian Mandible,” Ann. S. Afr. Mus., vol. xxviii, pt. ii, 1930. 
Hauvcuton, 8. H., and Boonstra, L. D.—*‘ Pareiasaurian Studies. Part VI.—The 
Osteology and Myology of the Locomotor Apparatus.—A. Hind Limb,” 

Ann. 8. Afr. Mus., vol. xxviii, pt. ili, 1930. 

Newton, E. T.—‘On Some New Reptiles from the Elgin Sandstones,” Phil. 
Trans. Roy. Soc. (Lond.), B, vol. clxxxiv, pp. 473-489, 1893. 

OweEN, R.—‘‘Catalogue of the Fossil Reptilia of South Africa, etc.,’’ Publ. British 
Museum, London, 1876. 

SEELEY, H. G.—“‘ Researches on the Structure, Organization, and Classification of 
the Fossil Reptilia. I1.—On Pareiasaurus bombidens (Owen), etc.,” Phil. 
Trans Roy. Soc. (Lond.), B, vol. clxxix, 1888. 

SEELEY, H. G.—‘“‘ Researches on the Structure, Organization, and Classification 
of the Fossil Reptilia. VII.—Further Observations on Pareiasaurus,” Phil. 
Trans. Roy. Soc. (Lond.), B, vol. clxxxiii, 1892. 

SusHxin, P. P.—‘On the Modifications of the Mandibular and Hyoid Arches and 
their Relations to the Brain-case in the Early Tetrapoda,” Pal. Zeitschrift, 
Bd. viii, Heft 4, 1927. 

VERSLUYS, J.—‘ Das Streptostylie-Problem und die Bewegungen im Schadel bei 
Sauropsiden,” Zool. Jahrb. Suppl., xv, pp. 545-716, 1912. 

Watson, D. M. S.—*‘ On the Skull of a Pareiasaurian Reptile, and on the Relation- 
ship of that Type,” P.Z.S., vol. i, 1914. 

Watson, D. M. S.—‘‘On the Nomenclature of the South African Pareiasaurians,” 
Ann. Mag. Nat. Hist., ser. viii, vol. xiv, 1914. 


Fig. 


EXPLANATION OF PLATES. 
Puate I.—Pareiasaur Skulls in Lateral View. x i. 


1. Bradysaurus baini. This is drawn from the type in the British Museum, 
but was checked and augmented by reference to South African Museum 
specimens (Nos. 3533, 4347, 4999, 5002, 5127, 9104, 9105), and the specimen 
in the Tiibingen Museum. 


. 2. Bradysaurus seeleyi. Drawn from the type and co-type in the British 


Museum, but checked and elaborated by reference to 8.A.M., Nos. 5624, 9115, 
9121, 9137, and 9168. 


. 3. Bradysaurus vanderbyli. Drawn from the type in the South African 


Museum (No. 3718) and two other specimens (Nos. 8941 and 9169). 


. 4. Bradysuchus whaitsi. Drawn from Broom’s published photograph of the 


type in the American Museum (No. 5567). 


. 5. Nochelesaurus alexandert. Drawn from the type in the South African 


Museum (No. 6239). 


. 6. Nochelesaurus strubenit. Drawn from a South African Museum specimen 


(No. 5019). 


. 7. Dolichopareia angusta. Drawn from the type in the South African Museum 


(No. 6238), but checked by reference to No. 3717. 


. 8. Brachypareia rogersi. Drawn from the neo-type in the South African 


Museum (No. 5012) and checked by No. 5340. 


g. 9. Brachypareia watsoni. Drawn from the type in the South African Museum 


(No. 6240). 


. 10. Embrithosaurus schwarzi. Drawn from the type in the South African 


Museum (No. 8034) and incorporating some details elucidated by Watson in 
the Hottentots River specimen. 


g. 11. Anthodon serrarius. Drawn from the South African Museum specimens 


(Nos. 4020 and 10074) and incorporating the details elucidated by Watson 
in the type in the British Museum. 


. 12. Pareiasuchus peringueyi. Drawn from the type in the South African 


Museum (No. 2337), with additional details from No. 2367. 


g. 13. Pareiasuchus nasicornis. Drawn from the type in the South African 


Museum (No. 3016). 


. 14. Pareiasaurus serridens. Drawn from the neo-type in the Geological 


Museum of the University of Stellenbosch. 


. 15. Scutosaurus karpinskii. Drawn from the published photograph by 


Hartmann-Weinberg. 


. 16. ‘ Pareiasaurus”’ horridus. Drawn from a cast in the British Museum. 
_17. Elginia mirabilis. Drawn from the published illustration of Newton. 


Newton’s determination of the sutures are doubtful. 


ir —troneale P. Orb. =postorbital. 
Jug. =jugal. Pr. Fr. =prefrontal. 
Lac. =lacrymal. Pr. Mx. =premaxilla. 
Mx. =maxilla. Qu. Jug. =quadratojugal. 
Na. =nasal. Sq. =squamosal. 

Pa. =parietal. Tab. =tabular. 


P. Fr. = postfrontal. 


i 
| Plato 1. 


Ann. 8. Afr. Mus., Vol. XXXT. 


Bia, 7. 


Fig. 13. Fro. 16, 


Neill & C0, Ly 


Fo. 3. 


eR Cs 


tie 


ANIA Raye! Payty Fy 


: 


a a er ee ee ee 


For details of specimens examined see explanation to Plate I. 


. 1. Bradysaurus baint. 

. 2. Bradysaurus seeleyi. 

. 3. Bradysaurus vanderbyli. 
. 4. Bradysuchus whaitsi. 
and a photograph by the American Museum. 


. Nochelesaurus alexanderi. 


. Nochelesaurus strubent. 


. Brachypareia rogersi. 


5 
6 
. 7. Dolichopareia angusta. 
8 
9 


. Brachypareia watsoni. 


. 10. Embrithosaurus schwarzi. 
. Ll. Anthodon serrarius. 

. 12. Pareiasaurus serridens. 

. 13. Pareiasuchus peringueyt. 
. 14. Pareiasuchus nasicornis. 

eg. 15. “‘Pareiasuchus”’ horridus. 
g. 16. Elginia mirabilis. 


Puate I1.—Pareiasaur Skulls in Dorsal View. x 


Drawn from a sketch kindly supplied by Dr. Broom 


Newton’s determination of the sutures are probably 


incorrect. 
Fr. =frontal. P. Orb. =postorbital. 
Jug. =jugal. Pr. Fr. =prefrontal. 
Lac. =lacrymal. Pr. Mx. =premaxilla. 
Mx. =maxilla. P. Pa. =postparietal. 
Na. =nasal. Qu. Jug. =quadratojugal. 
Pa. =parietal. Sq. =squamosal. 


P. Fr. = postfrontal. Tab. =tabular. 


Fia. ll. 


eo 

yeah \ 
SecccZ 

Tae is | 


SIN 
\ 
wy 


crate NTI Tak 
MO Lor A 
AONE, 


~ Ann. 8. Afr. Mus., Vol/ XN XT. 


Plato If. 


HGH 


i, 


Fig, 11. 


Fro. 16. 


Neill & 00, ld, 


oon oa»ark WH 


| 
=) 


ee 
Hm CW bo 


PuateE III.—Pareiasaur Skulls in Ventral View. x }. 


For details of specimens examined see explanation to Plate I. 


. Bradysaurus baini. 

. Bradysaurus seeleyi. 

. Bradysaurus vanderbyli. 

. Nochelesaurus strubent. 

. Brachypareia rogersi. 

. Embrithosaurus schwarzi. 

. Anthodon serrarius. 

. Pareiasuchus peringueyi. 

. Pareiasuchus nasicornis. 

. Pareiasaurus serridens. 

. Scutosaurus karpinskii. Details published by Sushkin are incorporated. 
. Koalemasaurus acutirostris. Drawn from the type in the Albany Museum. 
. Dolichopareia angusta. 

. Dolichopareia angusta. Dorsal view of the palate of the type (S.A.M., 
No. 


6238). 
A. Pter. =anterior ramus of the pterygoid. 
B. Oe: = basioccipital. 
B. Sph. = basisphenoid. 
For. =foramina in the premaxillary processes. 
For. Qu. =foramen quadrati. 
Int. Pter. Shit. =interpterygoid slit. 
Jug. =jugal. 
lis ter. =lateral ramus of the pterygoid. 
Mix: = maxilla. 
Pal. = palatine. 
Pe Oe: = paroccipital. 
P. Sph. = parasphenoidal rostrum. 
Pear: = postparietal. 
Pre He: = prefrontal flange. 
Prix: = premaxilla. 
Pre V0: = prevomer. 
Ree Ot: = pro-otic. 
Pe Pter. = posterior ramus of the pterygoid. 
Qu. =quadrate. 
Qu. Jug. = quadratojugal. 
Sub. Orb. Vac. =suborbital vacuity. 
SOc =supraoccipital. 
Sq. =squamosal. 
Tab. =tabular. 
Trans. =transversum. 


CASt: =tubercle for attachment of stapes. 


2 


50 c0Soc0~- ot 


Ann, 8. Afr. Mus., Vol. XXXT. 
3 ‘Plate U1. 


Boe ce 


[sooo 
Si igecee 


=a 


Neill & Co, Mids 


Puate IV.—Pareiasaur Skulls in Occipital View. x i. 


For details of specimens examined see explanation to Plate I. 


Fig. 1. Bradysaurus baini. 

Fig. 2. Bradysaurus seeleyi. 

Fig. 3. Bradysaurus vanderbyli. 

Fig. 4. Nochelesaurus alexanderi. 

Fig. 5. Nochelesaurus strubent. 

Fig. 6. Dolichopareia angusta. 

Fig. 7. Brachypareia waisoni. 

Fig. 8. Embrithosaurus schwarzi. 

Fig. 9. Pareiasuchus peringueyi (No. 2337). 

Fig. 10. — Be (No. 2367). After Haughton. 
Fig. 11. Scutosaurus karpinskii. After Hartmann-Weinberg and Sushkin. 


g. 12. Pareiasaurus serridens. From Annals of the University of Stellenbosch. 


B. Oc. —=basioccipital. 

B. Sph. =basisphenoid. 

Ex. Oc. =exoccipital. 

For. Mag. =foramen magnum. 

For. Qu. =foramen quadrati. 

P. Oc. =paroccipital. 

ED: = postparietal. 

P. Pter. =posterior ramus of the pterygoid. 
P.T.F. |=post-temporal fenestra. 
Qu. = quadrate. 

Qu. Jug. =quadratojugal. 

Sc: =supraoccipital. 

Sq. =squamosal. 


Tab. =tabular. 


. S. Afr. Mus., Plate IV.‘ 


Neill & Co., Lid, 


> 
Ann, 8, Afr. Mus., Vol. XXXT. Plato IV. * 


Neill & Co., Ltd, 


Fig. 
Fig. 
Fig. 
Fig. 
Fig. 
Fig. 
Fig. 
Fig. 
Fig. 
Fig. 
Fig. 


fo) 


Fig. 
Fig. 
Fig. 
Fig. 


oe owtn or WN 


— 
= © 


i eo 
Hq GW bo 


C 


— 
oO 


. Bradysaurus baini. 5th left tooth. 

. Bradysaurus seeleyi. 1st left tooth. 

. Bradysaurus vanderbyli. 3rd right tooth. 
. Nochelesaurus alexanderi. 6th left tooth. 
. Nochelesaurus strubeni. 5th left (mandibular) tooth (after Broom). 

. Dolichopareia angusta. 8th left tooth. 

. Brachypareia rogersi. 8th right tooth. 

. Brachypareia watsoni. 14th right tooth. 

. Embrithosaurus schwarz. 9th right tooth. 

. Anthodon serrarius. 6th or 7th left tooth (after Owen). 

. Propappus omocratus. 3rd or 4th right tooth. 

. Pareiasuchus peringueyi. 5th right tooth. 

. Pareiasuchus nasicornis. 4th right tooth. 

. Pareiasaurus serridens. 6th right tooth. 

. “ Pareiasaurus” russouwt. 3rd right mandibular tooth (from Haughton). 


Puate V.—Pareiasaurian Teeth in External View. x 3. 


Ann. S. Afr. Mus., Vol. XX XI, Plate V. 


r ha ney Sve 
cf [fy \ 
Au ti \, al 
Fic. 6. nie! 72 Fic. 8. Fig. 9. Fic. 10. 


| lh | te 
y 
\ i : i 5 
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\ Nh 
Wiad ql ul Wy 


Fic. 11. Pie. 12: iG. Fic. 14. 1 acepee ays 


Neill d& Co., Lid. 


VOL. XXXI, PART 1. 4 


Pirate VI. 


a Bradysaurus baini .  —. +~S.A.M., No. 50 


; (a) Lateral view. ‘ oe 
| ee (b) Dorsal view. 
| . 


2, aie 
m1 


i eee eee eee 


Ann. S. Afr. Mus., Vol. XX XI. 


Plate VI. 


NEU dP COn, iLial. 


nd 7 
a 5 
oe ; 
4%) 
a 
‘el 7 _ 
{2 eel 
a ie 
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of 
; ; 
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Pirate VII. 


S.A.M., No. 9137. 


. 


(a) Lateral view. 


Bradysaurus seeleyi 


(6) Dorsal view. 


4 
§ 
’ 

; 
q 


Ann. S. Afr. Mus., Vol. XX XI. Plate VII. 


Neill & Co., Ltd 


ya a ne Sa To 


Puate VIII. 
' (a) Bradysaurus seeleyi eee : S.A.M., No. 9137. 
(b) Bradysaurus vanderbyli S.A.M., No. 3718. 


= 


Ann. S. Afr. Mus., Vol. XX XI. 


Elate. Vil 


Neill & Co., Lid. 


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: é = 
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Puate IX. 


view. 


Bradysaurus vanderbyli 
(a) Lateral 
(6) Dorsal view. 


Ann. S. Afr. Mus., Vol. XX XI. Plate IX. 


Neill & Co., Ltd. 


Ue ey 
ne 


A 
ia 


Photograph 
merican 


view. 


Bradysuchus whaitsi . 
Lateral 


t a 
 —_ a iS ee 
i iy ies | iene eee ag aha ms 


y- = ; - ' i F 
Ve aces 8S EF IS Soe ga Se ee ee ee 


aa a mit = arc Sts - —— Se ai 
+= —————_———————————————— eee ——— ——_____—. — SS Ee ee ee 


Plate X. 


Ann. S. Afr. Mus., Vol. XX XI. 


va 


Puate XI. 


\ 
i 


No. 6239 


S-AM 


ere 


Nochelesaurus alexand 


(a) Lateral view. 
(b) Dorsal view. 


Plate XI. 


Ann. S. Afr. Mus., Vol. XX XI. 


Neill & Co., Ltd. 


Rn 
< 


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1 

24 

, 
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a 
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n 


Nochelesaurus strubeni ‘ 
; (a) Lateral view. 


(6) Dorsal view. 


Prats XI. 


S.A.M., No. 5¢ 


Ann. S. Afr. Mus., Vol. XX XI. 


Plate XII. 


Neill & Co., Lid. 


sae 


ee 


a > OA Se eee 


Puate XIII. 


(a) Nochelesaurus strubeni . S.A.M., No. 5019. 
(b) Dolichopareia angusta . S.A.M., No. 6238. 
¢ of 


Ann. S. Afr. Mus., Vol. XXXI, Plate XIII. 


S 


Neill & Co., Ltd. 


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LATE XIV. 


| Dolichopareia angusta . + S.A.M., No. 
(a) Dorsal view. - 


(6) Ventral view. 


Plate XIV. 


Ann. S. Afr. Mus., Vol. XX XI. 


Neill & Co., Ltd, 


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Brachypareia rogersi Pee eo 
(a) Lateral view. . SAM, No, 5012.amm 
(b) Lateral view. . §S.A.M., No. 4350. 
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Ann. S. Afr. Mus., Vol. 


XXXI. 


Plate XY. 


Neill d& Co., Ltd. 


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PLATE XVI. 


Brachypareia watsoni . . S.A.M., No. 6240. 
(a) Lateral view. 


(b) Dorsal view. 


- 

, 

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| 

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7 ad 
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Ann. S. Afr. Mus., Vol. XX XI. 


Plate XOV I. 


Neill & Co., Lid. 


sie isees M A aAll re v 
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Ys , we 
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PLATE XVII. 
Embrithosaurus ge pnne . “S.AM, No: 
(a) Lateral view. ig) Sse 


(6) Occipital view. . x }.- 


1 
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. 
1 
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™rk 
“s 
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: 
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Ann. S. Afr. Mus., Vol. XX XI. 


Plate XVII. 


Neill & Co., Ltd, 


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XXXI, PART 1. 7 
oy . % 
r¢ - : 4 ; 


Puate XVIII. 
re Pareiasuchus peringueyi . S.A.M., No. 2337. « 


ea , (a) Lateral view. 


: 
- 
’ 


(6) Occipital view. 


= 


i ei eee a ee ee ee ES 


Ann. S. Afr. Mus., Vol. XX XI. Plate XVIII. 


Neill & Co., Lid. 


nile Aas r 7 
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| 
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PLATE XIX. 


Pareiasuchus peringueyi . S.A.M., No. 2337. 
(a) Dorsal view. 


_ (b) Ventral view. 


. a ae 
oe —= Ais 
hit ate i 


Ann. S. Afr. Mus., Vol. XX XI. Plate XIX. 


eill & Co., Ltd. 


SSS 


Poke "s Boy 
Lee ae ey me 
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PS Rae ae 
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ie Puate XX. ' 


Pareiasuchus nasicornis . S8S.A.M., No. 3016. 


(a) Ventral view. 


(6) Dorsal view. 


Ann. 


S. Afr. Mus., Vol. XX XI. 


Plate XX. 


Neill & Co., Ltd. 


= : 
= 
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Prarin XOX, 


(a) Pareiasuchus nasicornis §.A.M., No. 3016. Lateral view. 


| (b) Anthodon serrarius . §.A.M., No. 10074. Lateral view. 


‘ 


Ann. 


S. Afr. Mus., Vol. XX XI. 


Plate X XI. 


Neill & Co., Lid. 


PuLaTE XXII. 


(a) Anthodon serrarius . Amer. Mus., No. 7001. Dorsal view. x 4. 
Photograph kindly supplied by the 
American Museum. 


(b) Pareiasaurus serridens . Geol. Mus. Univ. Stellenbosch. Lateral 
view. x #. 


Plate XXII. 


Amn. S. Afr. Mus., Vol. XXXI. 


i 


S 


Neill & Co., Ltd. 


(30m) 


Pareiasaurian Studies. 


Part X.—The Dermal Armour.—By Likuwr D. Boonstra, D.Sc., 
Department of Palaeontology, South African Museum. 


(With Plates XXITI-XX VII.) 


INTRODUCTION. 


THat the Pareiasaurs possessed a more or less extensive armour of 
bony scutes of dermal origin has been noted by a number of authors. 
Seeley gave the first and also the most complete account of the 
armature in the genus Bradysaurus. Broom described it in Pro- 
pappus and Brachypareia, and referred to it in Embrithosaurus and 
Anthodon; publishing jointly with Haughton, a brief reference was 
made to the scutes of Parevasuchus. Subsequently Watson used the 
nature of the armature as a generic character. Amalitzky and 
Hartmann-Weinberg have considered the nature of the scutes as of 
specific as well as of generic value in the case of the Russian 
Parelasaurs. 

In the body of this paper an attempt is made to gather together our 
knowledge of the Pareiasaurian armour in order to obtain a general 
idea of the appearance and the defensive abilities of the Pareiasaurs 
as a whole, and, secondly, to determine the specific and generic 
differences (if any), and, as a further development of the same theme, 
to see whether there is any progressive or retrogressive development 
of the armature. The large number of new specimens now in the 
South African Museum, together with those already partly known, 
fully justifies such an attempt being made. Wherever possible the 
verbal description is accompanied by photographs. 


Genus BRADYSAURUS. 


1888. Seeley, H. G., Phil. Trans. Roy. Soc., vol. clxxix, B 22, p. 91. 
1892. Seeley, H. G., Phil. Trans. Roy. Soc., vol. clxxxii, B 76, 
pp. 345, 346. 


40 Annals of the South African Museum. 


1908. Seeley, H. G., P.Z.8., vol. 1, pp. 605-607. 
1914. Watson, D. M.8., Ann. Mag. Nat. Hist., vol. xiv. 


The dermal ossicles in the three species of this genus are very similar 
(Pl. XXIII; Pl. XXIV, fig. 1); the scutes preserved are roughly 
circular or oval in outline; the external surface is relatively smooth; 
in Br. vanderbyli they are practically devoid of any ornamentation; 
in Br. baini the scutes are convex, curving down from a thickened 
centre which in some scutes appears like an incipient boss; there are 
no distinct radiating ridges; in Br. seeleyr the central boss appears 
to be somewhat more distinct, and the surface somewhat pitted. 
There is also very little difference in the size of the scutes, viz. Br. 
baini (50 x 50-45 x12 mm.), Br. seeleys (57-35 x 50-30 x 10-7 mm.), 
Br. vanderbyli (48 x 47 x 7 mm..). 

The above facts appear to exhaust the dissimilarities. From the 
available facts the following composite account, which includes 
some of Seeley’s observations, of the arrangement of the armour 


appears to be correct for all three species. “No indications of large 
dermal bones are preserved. Three rows of scutes appear to have 
extended down the median line of the back.” “The median row was 


placed on the summits of the neural spines and above the interspaces 
between them. In contact with these, laterally, there is a pair of 
scutes extending transversely outward; so that in the antero-posterior 
direction three scutes may be counted corresponding to each vertebra. 
And the effect is a median longitudinal strip of close-set scutes, 
flanked by two lateral rows.” From the large number of specimens 
in the South African Museum it appears that the scutes were in more 
close contact above the scapulae and the sacrum than along the rest 
of the back and on the tail. Above the ribs, 7.e. on the flanks of the 
animal, the scutes seem to have been little developed, and this also 
applies to the upper portions of the limbs. No small stud-shaped or 
conical ossicles have been found in this genus, and they probably 
did not exist. 

In this connection it is perhaps significant to note that in the other 
large genera (Dolichopareia and Nochelesaurus) of the Tapinocephalus 
zone no scutes have been found. It must be admitted that of these 
genera we only know a few specimens, not very fully preserved, but 
the absence of scutes in the specimens known seems to indicate a 
lesser development of armour than in the more highly evolved forms 
of the younger horizons. 


Pareiasaurian Studies.—Part X. 4] 


Genus KOALEMASAURUS. 


No dermal scutes are preserved in association with the type skull 
in the Albany Museum. Associated with a skull and vertebral 
column referred by me to this genus there are, fortunately, a few 
scutes. Although it is impossible to ascertain the extent of the 
armature from this scanty material, the nature of an individual 
scute can be described. The scutes preserved (Pl. XXIV, fig. 2) are 
small (45 x32x7 mm.), oval in outline, smooth, with only a slight 
indication of a central knob and a hardly perceptible rugous radiation. 

The possibility exists that the scantiness of the number of scutes 
preserved is due to the fact that the specimen is immature, and to 
the same cause the smoothness of the scutes may be ascribed. 


Genus BRACHYPAREIA. 


ie broom. kh. Anns, Air. Mus, vol. vii, p. 323, 
1914. Watson, D. M.8., Ann. Mag. Nat. Hist., vol. xiv. 


Unfortunately no scutes are known of Brachypareia watsoni—only 
the skull of the type specimen being preserved. The description that 
follows thus only refers to Brachypareia rogersi. A very large number 
of dermal scutes of various sizes and shapes are preserved in associa- 
tion with the type specimen (PI. X XIV, fig. 3). There is a perfect 
gradation from fairly large ovoid or circular ossicles, whose outer 
surfaces are ornamented with a central knob from which fairly strong 
ridges radiate, through a series of smaller circular ossicles with a 
strong central knob, but with only feeble ridges radiating from it, 
standing on a base and presenting the appearance of a depressed 
collar stud, to small globular ossicles about the size of large peas, and 
with apparently no surface ornamentation. In all the ossicles, with 
the exception of the globular ones, there is on the external surface a 
foramen entering the interior of the bone and cut into the side of the 
knob. It communicates with a small foramen opening on the internal 
surface. From the plate a better idea can be obtained of the relative 
sizes and shapes than from any verbal description. 

The number of ossicles preserved, in addition to this gradation in 
a series in point of size and ornamentation, allows one to draw the 
following conclusions, viz. that all along the back, from the neck to 
the proximal dorsal surface of the tail, there must have been rows of 
ossicles (two or three per rump segment) with contiguous edges. 
The flanks, withers, and quarters were apparently studded with the 


42 Annals of the South African Museum. 


circular stud-shaped ossicles; the upper portions of the limbs, the 
extremity of the tail, and the lateral parts of the belly must have 
housed the smaller globular ossicles; the belly, throat, and those 
parts affected by the movements of the limbs would have been 
practically free of dermal armour. 


Genus EMBRITHOSAURUS. 


1903. Broom, R., Ann. 8. Afr. Mus., vol. iv, pl. xvi. 
1914. Watson, D. M.S., Ann. Mag. Nat. Hist., vol. xiv. 


Although Broom only mentioned the dermal ossicles of Embritho- 
saurus in his explanation to a plate, a series, above the 6th, 7th, and 
8th presacral vertebrae, is very well preserved in situ and merits a 
fuller description. 

Above the spinous process a large (50 x 50 mm.) and thick (20 mm.) 
scute is situated (Pl. XXYV, fig. 1); it has a very large and strong 
central boss of irregular shape, but no ridges radiate from it towards 
the periphery. Both anteriorly and posteriorly this scute articulates 
with a smaller smoother scute, which overlies the interspace between 
the spinous processes; laterally there lie two rows of scutes, each 
row has two scutes per segment—one lying on the postzygapophysial 
ridge and the other in the interspace; these scutes articulate with 
each other and with the median row of scutes. These lateral scutes 
are smaller (50 x 40 mm.) and thinner (15 mm.), have a more circular 
boss from which a few ridges commence, but do not reach the 
periphery; theedgesareserrate. Astillsmaller scute (35 x 22 x5 mm.), 
apparently from the flank, is preserved. It is oval in outline, has a 
slightly thickened centre, rugose near the edges, and serrated edge. 

In summary one can thus state that above the withers of the 
animal there were five rows of articulating ossicles, that it is probable 
that some of these rows extended along the whole length of the back, 
and that the flanks were in all probability studded with smaller and 
smoother scutes. 


Genus ANTHODON. 


1930. Broom, R., Amer. Mus. Novitates, No. 448. 


Although Broom has described the specimen of Anthodon in the 
American Museum of Natural History as a new species, I have seen 
fit to doubt that it is distinct from Owen’s species. The nature of 
the dermal armour in a specimen in the South African Museum, which 


Pareiasaurian Studies.—Part X. 43 


certainly belongs to Owen’s species, is so similar to that of the 
American Museum specimen that my doubt as to the validity of 
Broom’s species appears to have further justification. 

Broom has stated that the back of Anthodon was “covered by a 
bony carapace formed by large articulating bony scutes.”’ From a 
number of dermal ossicles found overlying the sacral region of one of 
the South African Museum specimens, it becomes evident (Pl. XXV, 
fig. 2; Pl. XXVI, fig. 1) that at least some of the scutes are firmly 
articulated to each other, so that in ventral view (Pl. X XVI, fig. 1) 
the line of articulation is only indicated by a digitating suture similar 
to those found between some cranial bones. A small foramen is 
also to be noted opening on to the ventral surface. 

The scutes are of medium size (64-54 x 45-40 x 11 mm.), roughly 
circular in outline, and are strongly ornamented on the outer surfaces, 
the central knob stands out very distinctly—this being due to the 
fact that it does not grade into the base, but is partly demarcated 
from it by an encircling moat traversed by radiating ridges. Relative 
to this moat the peripheral edge is somewhat raised and thickened; 
at the articulation with adjoining scutes the edge is deeply serrate. 


Genus PAREIASAURUS. 


1914. Watson, D. M.8., Ann. Mag. Nat. Hist., vol. xiv. 


Watson has shown that the pelvis described by Owen as that of 
Dicynodon tigriceps belongs to the type of Pareiasaurus serridens. 
With it are associated scutes which would appear to have formed an 
extensive armour. “The whole of the dorsal surface of the pelvis 
between the crests of the ilia is covered with very large scales which 
are of the Propappus type, but differ in their more definitely pitted 
ornament and much larger size.” Associated with the neo-type 
skull in the University of Stellenbosch there are a number of scutes 
preserved, and I also have a specimen in the South African Museum 
with a number of scutes (Pl. X XVI, fig. 2). 

Three sizes (67-53-47 x 58-45-40 x 25-20-15 mm.) of ossicles are 
preserved. Each ossicle has a strong central boss from which strong 
ridges alternating with deep grooves radiate towards the periphery. 
The foramen, entering the interior of the bone on one side of the boss, 
is constant. A small foramen also opens on the internal surface of 
the scute. The scutes of Pareiasaurus are very similar to those of 
Parevasuchus. 

There is evidence (Pl. X XVI, left bottom pair) that some of the 


44 Annals of the South African Museum. 


lateral scutes overlying the ribs have their serrated edges interlocking, 
and if this is the case in a series of scutes a carapace of mosaic pattern 
would be formed, but it does not appear that a very great degree of 
rigidity would result as there appears to be a possibility of relative 
movement at the “sutures.” 

The scutes overlying the ribs have their longer diameter directed 
dorso-ventrally and the shorter antero-posteriorly. There is, further- 
more, reason to believe that, as in the case of the dorso-median scutes, 
the lateral scutes on the flanks were also regularly arranged in rows. 


Genus PAREIASUCHUS. 


1913. Broom, R., and Haughton, 8. H., Ann. 8. Afr. Mus., vol. xu, 
p- 25. 


Dermal scutes are present in the type specimens of both species. 
Those of Pareiasuchus peringueyt have been described by Broom 
and Haughton in the following words: ‘Dermal ossicles occur all 
along the back, between the proximal parts of the ribs, especially 
in the neck, and around the pelvis. The scutes are smaller than 
those of Propappus.” Those of P. nasicornis have as yet not been 
described. Although only a limited number of bony scutes are 
preserved there is sufficient evidence available to enable one to state 
that fairly large (65-40 x 55-30 x 15-10 mm.) scutes were present, in 
at least three rows, all along the back, and that they appear to have 
been especially well-developed above the shoulder-blades and the 
sacrum; as preserved, some of the scutes are in contact along their 
edges, and it is thus legitimate to assume that the armour consisted 
of rows of scutes in contact with each other. As is apparent from the 
illustration (Pl. XX VII, fig. 1), the scutes vary in size, shape, and ex- 
ternal ornamentation. All the preserved scutes are rugose; they have 
a central knob from which strong ridges radiate; the foramen on the 
side of the knob is constant, but in one case the foramen is open to 
the periphery and forms a notch. 


Genus PROPAPPUS. 


1908. Broom, R., Ann. 8. Afr. Mus., vol. iv, pp. 358, 359. 
1914. Watson, D., M.S. Ann. Mag. Nat. Hist., vol. xiv. 


Although only a few scutes of Propappus parvus are preserved in 
the one known specimen, it appears that the armour differs sub- 
stantially from that of Propappus omocratus. 


Pareivasaurian Studies.—Part X. 45 


The armour of Propappus omocratus was first described by Broom : 
“Propappus must have had the whole back and probably sides 
covered with a carapace of bony plates somewhat after the manner 
of the crocodile. They vary in size from plates 50 mm. in diameter 
to little bony nodules about the size of peas. The larger ones have a 
central boss and irregular radiating ridges. At the edges they overlap 
each other, and it seems probable that the carapace was not quite 
rigid. It extended at least as far out on the sides as 300 mm. from 
the middle line. There is no evidence of any plastron.”’ 

Some additional remarks may be appended (Pl. XXVII, fig. 2): 
there is evidence that along the median line in each rump-segment 
one scute overlay the spinous process, one lay between the spinous 
processes, and lateral to these one lay on the postzygapophysial ridge, 
and one in the interspace between this and the next ridge. Broom 
records an overlapping of the scutes, whereas to me there appears 
to be an articulation at the edges. 

Only a few scutes of Propappus parvus are preserved (Pl. X XVII, 
fig. 2). Judging from these the scutes appear to be more oval (60-50 x 
37 mm.) and flatter (8 mm.) than those of Pr. omocratus; the central 
boss is weaker and the ornamentation much weaker—the radiating 
ridges being only distinct at the edges of the scute. 


Genus ScUTOSAURUS. 


1922. Amalitzky, V., Bull. Acad. Sc. Russ. 
1930. Hartmann-Weinberg, Pal. Zeitschr., Bd. xi. 


In 1922 Amalitzky, in his brief ‘“‘diagnoses,” mentioned the dermal 
armour of the Russian Pareiasaurs. In Scutosaurus Karpinski, 
“star-shaped dermal plates are situated above the spinous processes 
of the vertebrae, two or three rows of similar plates lie on the sides 
of the body. There are also plates distributed, without apparent 
order, over the whole body (fig. 8) (from the illustration these 
ossicles look very similar to those of Parevasaurus L.D.B.); the belly 
was covered with small conical bosses.” 

“ Pareiosaurus”’ tuberculatus has “three rows of star-shaped dermal 
plates on the sides of the neck; in the tail region the plates are 
boss-like.”’ : 

In 1930 Hartmann-Weinberg stated that the osseous scutes of 
Scutosaurus are of various shapes and sizes. The ornamentation is 
similar to that of the skull; they lie closely applied to each other to 
form a heavy carapace. 


46 Annals of the South African Museum. 


Incerta sedis. 


1908. Seeley, H. G., P.Z.S., pp. 607-610. 


In 1908 Seeley described and figured (text-figs. 125 and 126) a 
number of scutes which he had obtained from Steenkamps Poort. 
The evidence afforded by the nature of these scutes appeared to him 
to justify the creation of a new species, Parevasaurus steenkampensis. 
From his excellent description and figures it appears that these scutes 
are very similar to those of either Brachypareia rogersi or Pareiasuchus 
nasicornis, but it would be rash, lacking corroborative evidence from 
the rest of the skeleton, definitely to assign these scutes to either of 
these forms. These scutes are thus best referred to as simply Pareia- 
saurian, with the corollary that they definitely do not belong to forms 
of the genus Bradysaurus. 


GENERAL. 


It is possible, from the above detailed account, to determine the 
essential characters of the Pareiasaurian dermal armour. One of the 
most striking features is the apparent correlation that exists between 
the ornamentation on the dermal cranial bones and that on the 
dermal ossicles. Most of the dermal cranial bones have a more or 
less centrally placed boss from which ridges and grooves and bone- 
fibres radiate, and the sutures between adjoining bones is often of a 
zigzag nature. Practically the same features are found in the dermal 
ossicles. 

Moreover, in the early forms with moderately smooth skulls the 
dermal ossicles are found to be fairly smooth, and conversely, the 
rugose skulls of later forms are accompanied by highly ornamented 
and closely articulating dermal ossicles. It thus appears to be con- 
sistent with the facts to conclude that the Pareiasaurs during their 
evolutionary history, in addition to acquiring a greater cranial 
rugosity, developed an armour of more highly ornamented and more 
closely articulating dermal ossicles. 

It would, furthermore, appear that the development of the dermal 
armour began above the scapulae and above the sacrum, and then 
extended along the whole back; then along the flanks, and finally 
above the limbs and possibly also the belly. So also the articulation 
of the scutes, inter se, appears to have followed the same order of 
development. 

There seems to be little doubt that in general the dermal ossicles 


Pareiasaurian Studies.—Part X. 47 


were arranged in regular rows running parallel and at right angles 
to the dorsal median line. The arrangement of a pair of ossicles per 
vertebral segment and also the position in relation to the ribs seems 
to indicate that the scutes were more firmly articulated along their 
lateral margins than along either the anterior or posterior margin, 
so that it is perhaps more correct to refer to transverse rows of scutes. 
I imagine the arrangement of the scutes to have been very similar 
to those of the living Zonurus. 

Mention has been made of a foramen entering each ossicle on the 
external surface on the side of the central bess, and also of a much 
smaller foramen opening on the internal surface. The former opens 
into a conical pit on the outer surface. In specially prepared sections 
it is seen that these foramina communicate; I have examined the 
scutes in the living Varanus and Zonurus, and find that in the former 
a nutritive foramen enters on the external surface, whereas in the 
latter it enters on the internal surface just under the spine. It would 
thus appear that both foramina in the Pareiasaur scute, in all 
probability, housed blood-vessels. 

Consisting of strong bony ossicles, the Pareiasaur armour was 
certainly of great biological value to these animals. As has been 
noted, it is first developed above the most vulnerable spots, and it is 
significant that the forms surviving up to the end of Lower Beaufort 
times show the highest development of armour. The last Pareia- 
saurs, although carrying a greater amount of armour, do not appear 
to have become more clumsy animals. On the contrary, our studies 
of the limbs and limb-musculature point to a more upright gait 
accompanied by greater agility. The development of armour in 
these animals was certainly an acquisition of a character of decided 
survival value, and, furthermore, does not appear to have eventually 
overreached itself. 


48 


Fig 


Fig. 
Fig. 


Fig. 


Fig. 


Fig. 


Fig. 


Annals of the South African Museum. 
EXPLANATION OF PLATES. 


Prate XXIII. 


. 1. Bradysaurus seeleyi (9168). x 3. Dermal ossicles above 5th, 6th, and 
7th presacral vertebrae. The median and first lateral row of each side 
are represented. 


. 2. Bradysaurus baini (9001). x 2. Dermal ossicles of the first lateral row 


above the anterior dorsal vertebrae. 


PratrH XOX Ly: 


. 1. Bradysaurus vanderbyli (9169). x 1. Three articulating ossicles above 
the anterior dorsal vertebrae. 

2. Koalemasaurus acutirostris (4345). x 3. Two isolated ossicles. 

3. Brachypareia rogerst (8453). x 2. <A series of isolated ossicles; the 
larger ones in the top row come from the median line of the back; the smaller 
ones in the second row from the flanks, and the lowest row probably low down 
the flanks and upper portions of the limbs. 


PLATE XXV. 


. 1. Embrithosaurus schwarzi (8034). x 3. Ossicles from the median and 


first lateral row of each side; from above the 6th, 7th, and 8th presacral 
vertebrae. 

. 2. Anthodon serrarius (10074). x 3. Ossicles of the median and first lateral 
row; from above the sacrum. 


PLATE XXVI. 


1. Anthodon serrarvus (10074). x 4. On the left is the external view of two 
closely articulating ossicles. On the right is the internal view of two ossicles 
very firmly joined—a suture marks the line of articulation. Both sets come 
from above the sacrum. 

2. Pareiasaurus serridens (10032). x 4. A series of isolated scutes; the 
two top ossicles probably came from the back; the left bottom pair are firmly 
articulated and overlie a rib and thus come from the flank—the long axes of 
these two scutes are at right angles to the long axis of the body; the two 
ossicles in the left bottom corner probably come from lower down the flank. 


Puate XXVIII. 


1. Pareiasuchus nasicornis (38016). x 2. A set of isolated scutes; the small 
scute with the notch is remarkable. 

2. Propappus parvus (2351). x 4. The two upper scutes belong to this 
species; the one on the left is seen in internal view, that on the right in external 
view; in the former a small foramen is to be noted. 

Propappus omocratus (1058). x 4. The ossicle on the bottom left corner 
is from the dorsal median row; that on the bottom right from a lateral row 
as it overlies a rib; smaller stud-shaped ossicles, similar to those of Brachy- 
pareia, are also preserved. 


Ann. 8S. Afr. Mus., Vol. XX XI. Plate X XIII. 


Neill & Co., Ltd. 


Ann. S. Afr. 


Mus., Vol. XX XI. 


ING, Be 


Plate XXIV. 


ie, Bs 


Neill d&: Co., Ltd. 


Ann. S. Afr. Mus., Vol. XX XI. Plate Oxeve 


I 


Fic. 


S 
Fe 


leill & Co., Ltd. 


= 
— ce ri 5 / 
: . 
| . ; : 
a ; 
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2 - . | 
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: 


Ann. S. Afr. Mus., Vol. XX XI. 


Fre. 2. 


Plate X XVI. 


Neill & Co., Lid. 


Ann. S. Afr. Mus., Vol. XX XI. Plate X XVII. 


Gale 


Nie, YD. 


Neill & Co., Ltd. 


Ves) 


Pareiasaurian Studies. 


Part X1.—The Vertebral Column and Ribs.—By Lizuwe D. BoonstRa, 
D.Sc., Department of Palaeontology, South African Museum. 


(With Plate XXVIII and Text-figures 1-7.) 


INTRODUCTION. 


To Owen (1876) we owe our first description (pp. 9-11) of a Pareia- 
saurian vertebra. But the vertebra figured (pl. xi) and described 
as that of Pareiasaurus bombidens is that of a Deinocephalian, in all 
probability Tapinocephalus, and not of a Pareiasaur at all. 

In 1888 Seeley described (pp. 77-89) and figured (pls. xii, xvii, 
Xvi, and xix) the nearly complete vertebral column of the Palmiet 
Fontein specimen of Bradysaurus seeleyi. According to this author 
there are 18 presacral, 2 anchylosed sacral, and 9 caudal vertebrae. 
The presacrals consisted of 9 cervicals and 9 dorsals. Only the first 
sacral rib supported the ium. There are 9 pairs of dorsal ribs. 
“The head of the rib is obviously notched.’ “In the last dorsal 
the rib appears to be almost entirely supported upon the diapophysis, 
the parapophysis having become very small.” 

In 1892 the same author described (pp. 338-345) and figured 
(pls. xvil, XxX1, xxl, and xxi) the nearly complete column of Brady- 
saurus bain. ““There appears to be eighteen presacral vertebrae.” 
Of these Seeley thought that 10 were cervicals and 8 dorsals. The 
sacrum consisted of 4 vertebrae—the first a sacro-lumbar, then a true 
sacral and 2 sacro-caudals. Four sacral ribs supported the iium— 
the first two being much more massive than the last two. “Twenty- 
six caudals are preserved”—but the tip of the tail was lost. The 
first 4 ribs had, according to Seeley, single facets; in the 5th to the 
7th the facet is elongated. From the 5th postsacral vertebra chevrons 
appear—the first being unpaired, the rest paired. 

In the Tamboer specimen of Bradysaurus seeleyr there were 19 
presacrals, of which 11 were dorsals and 8 cervicals. 

In 1893 Newton described (pp. 489-493) and figured (pl. xl1) parts 


50 Annals of the South African Museum. 


of the vertebral column of Elginia mirabilis. There are 4 anchylosed 
sacrals with ribs—the Ist being much larger and carrying a much 
stronger rib than the posterior three. 

Broom, in 1903, described (pp. 127-130) and figured (pl. xvi) the 
well-preserved vertebral column of Embrithosaurus schwarz. “There 
are 20 presacral vertebrae, 2 anchylosed sacrals, and probably 30 
caudal vertebrae.” Broom found that no rib articulated with the 
atlas; a two-headed rib articulated with the axis; in the 3rd, 4th, 
and 5th cervicals there were two articular surfaces for each rib; in 
the 6th the two articulatory surfaces became united, and in all the 
succeeding presacral vertebrae there is only a single articular surface; 
two sacral ribs give attachment to the ilium—the first is very much 
the stronger and there may have been a third sacral rib; chevrons 
were developed. 

The incomplete vertebral column of Propappus omocratus was 
described (pp. 353, 354) and figured (pl. xlv) by Broom in 1908, and 
that of Brachypareia rogersi (pp. 323, 324, pl. xix) in 1912. 

Broom and Haughton described (pp. 20-22) and figured (pl. v) the 
vertebral column of Parevasuchus peringueyi and found 19 presacrals, 
4 anchylosed sacrals, and 12 caudals; the axis had an apparently 
single-headed rib; the 3rd, 4th, and 5th cervicals had double-headed 
ribs; the 6th and all the other presacrals had single-headed ribs; 
there were 4 sacrals supporting the ilium—the lst was massive, the 
2nd and 3rd weak, and the 4th moderate; from the 7th caudal chevrons 
were present. 

An incomplete series of Propappus parvus was described (pp. 44-45) 
by Haughton. There were apparently 4 sacral ribs; the ribs were 
single-headed and there were chevrons on the caudal vertebrae. 

In 1914 Watson had found parts of the type of Parevasaurus 
serridens in the cellars of the British Museum. The sacrum consisted 
of 4 constricted vertebrae. These parts had been described (pp. 
40-41) and figured (pl. xxxvi, fig. 1) by Owen as belonging to Dicynodon 
tigriceps. Parts of Propappus omocratus were also found by Watson. 
The sacrum consisted of 4 vertebrae and was originally figured 
(pl. xxxvi, figs. 2-4 and figure on p. 74) by Owen as Dicynodon 
tigriceps and Dicynodon. Watson defines the genus Bradysaurus as 
having 4 sacrals, as also had Embrithosaurus. 

Haughton and Boonstra, in their paper on the Hind Limb, described 
and figured the various sacra and their ribs in so far as they acted as 
supports to the pelvic girdle. 

In her description of Scutosaurus, Hartmann-Weinberg stated: 


Pareiasaurian Studies.—Part XI. 51 


“Halswirbel 6, Rumfwirbel 13, Sacralwirbel 4-5, Schwanzwirbel 
genaue Zahl unbekannt.” 1-3 sacral ribs articulated with the 
ilium—3-2 probably ligamentously. The cervicals had two-headed 
and the dorsals single-headed ribs; the postsacrals were provided 
with chevrons. 


GENERAL OSTEOLOGY. 


Although there are in the collection of the South African Museum 
parts of the vertebral column of 46 Pareiasaurs, only a small propor- 
tion of these are even approximately complete. Quite a number 
have the dorsal series complete but very often lack a full series of 
cervicals, and in all, except one specimen, the postsacrals are not 
preserved as a complete series. 

In view of these facts it has been considered canbe to base our 
general description chiefly on the one column which is known to be 
practically complete and to use the other material mainly to check 
any conclusion arrived at. Some of this material belonging to the 
various species will, of course, be treated of in a subsequent section 
which deals with the vertebral column from a systemic aspect. 

The above-mentioned column (8.A.M., No. 9168) was excavated by 
the author on the farm Knoffelfontein in the Beaufort West Division 
and was that of a practically complete animal identified as Brady- 
saurus seeleyt. The skull was disarticulated and the atlantal centrum, 
intercentrum, and atlantal arch were not preserved. From the axis 
onwards, however, the column was absolutely complete—the posterior 
section of the tail was removed in a block of matrix which extended 
to the last postsacral vertebra so that there is no doubt as to the 
number of postsacrals. The atlantal centrum, intercentrum, and 
paired arch are, however, preserved in a specimen of Bradysaurus 
vanderbylr (S.A.M., No. 9169) found by the author at the same locality. 
These elements are in addition preserved in a specimen of Nochele- 
saurus strubent (S.A.M., No. 5590) from Abrahams Kraal, Prince Albert, 
and in a specimen of Pareiasaurus serridens (Stellenbosch, No. K. 218) 
from Houd Constant, Graaff-Reinet, and also, although much distorted, 
in a specimen of Nochelesaurus strubend (S.A.M., No. 5019) from 
Blaauwkrantz, Prince Albert, and, incompletely, in the type specimen 
of Brachypareia watson (S.A.M., No. 6240) from Abrahams Kraal, 
Prince Albert. 

The column consists of 20 presacrals, to which in all probability 
another, viz. the pro-atlas, should be added; 4 sacrals with coalesced 


52 Annals of the South African Museum. 


centra; and 39 postsacrals. These main divisions can be further 
subdivided, viz. 4 cervicals 15 dorsals, 4 sacrals, 3 pygals, and 
36 caudals. 

Cervicals (Pl. XXVIII, Text-figs. 1, 2, and 3).—Although the usual 
criterion for determining the beginning of the dorsal series, viz. the 


AxA 


TExt-FIG. 1.—Bradysaurus vanderbyli. S.A.M., No. 9169. 
Lateral view of the atlas and axis. x 1. 
At. A. =atlantal arch, left half. 
At. I. =atlantal intercentrum. 
Ax. A. =axiall arch. 
Ax. I. =axial intercentrum. 


C,. =atlantal pleurocentrum (odontoid). 

C,. =axial pleurocentrum. 

1D); = diapophyses on atlantal and axial arch. 

F. = foramen. 

Iles =intercentrum between the axis and the 3rd cervical. 

12 = parapophysial facets on the atlantal and axial intercentra. 
PZ. =axial postzygapophysis. 


sternum, is absent, there appears to be sufficient other evidence to 
enable one to state the number of cervicals. The first 5 presacrals 
carry two separate processes—a diapophysis on the arch and a para- 
pophysis on the centrum or on the preceding intercentrum; articu- 
lating with these there are 5 slender dichocephalous ribs which have 
the tuberculum and capitulum separated by a deep notch. The 
characters of the next vertebra—the 6th—show an abrupt change; 
the diapophysis and parapophysis are fused to form an elongated 


Pareiasaurian Studies.—Part XT. 53 


vertical transverse process, and articulating with this is a stout and 
much longer holocephalous rib. This abrupt change is considered 
to afford sufficient evidence for the delimitation of the cervical series. 

The lst vertebra differs from the rest in retaining certain primitive 
temnospondylous characters; the 2nd is specialised, as in all reptiles 
with the exception of Seymouria; the 3rd differs from the succeeding 
vertebrae in its distinctively shaped neural spine; the 4th and 5th 


TrExt-FIG. 2.—Hmbrithosaurus schwarzi. S.A.M., No. 8034. 
Anterior view of atlas and axis. x 4. 


The left half of the atlantal arch and half of the unpaired atlantal intercentrum 
have been omitted from the sketch. 


At. A. =atlantal arch. 
At. I. =atlantal intercentrum. 
Ax. A. =axial arch. 


C\. =atlantal pleurocentrum (odontoid). 

1D). = diapophyses on atlantal and axial arch. 
Pr. Z. =axial prezygapophysis. 

Z.? =an undeveloped zygosphene? 


have nearly all the characters of the dorsal vertebrae and pass 
gradually into the dorsal series. 

Proatlas.—In no specimen has any portion of a proatlas actually 
been found. The fact, however, that it occurs in all groups of 
primitive reptiles leads one to entertain the idea of its probable 
presence. Furthermore, there seems to be some evidence pointing 
that way, viz. the antero-dorsal extremity of the atlantal arch 
appears to be very similar to that in the forms possessing a proatlas 
(e.g. Ophiacodon and Moschops) and it seems highly probable that a 

vol, XXXI, PART I. 9 


54 Annals of the South African Museum. 


proatlas was articulated to it; in addition the relations of the occiput 
with the atlas is such that the dorsal surface of the medulla on 
emerging from the foramen magnum would be wholly uncovered in 
the absence of a proatlas. In forms like Scymnognathus and 
Aelurognathus this is precisely the function of the paired proatlas. 
For these reasons it 1s believed that a proatlas was present in life. 
Atlas (Pl. XXVIII, Text-figs. 1, 2, and 3).—The atlas, as in nearly 
all reptiles, remains temnospondylous; it consists of a paired neural 


AxA 


Ci 


Ata 


T3 Axl 


TExt-FIG. 3.—Bradysaurus vanderbyli. S.A.M., No. 9169. Median 
section through the atlas and axis. x 4. 
At. I. =atlantal intercentrum. 
Ax. A. =axial arch. 
Ax. I. =axial intercentrum. 


C,.  =atlantal pleurocentrum (odontoid). 
C,. =axial pleurocentrum. 
I,. =intercentrum between the axis and the 3rd cervical. 


arch resting in part on a large hemicylindrical intercentrum, and in 
part on an odontoid-like pleurocentrum; the whole is loosely arti- 
culated to the axis. The arch is a bone of complex shape; dorso- 
posteriorly it carries a process which corresponds to the processes 
of succeeding vertebrae that carry the downwardly facing post- 
zygapophyses ; it articulates with the prezygapophysis on the axis; 
dorso-anteriorly it probably gave articulation to a proatlas; ventrally 
it articulates with the intercentrum; ventro-posteriorly it carries a 
posteriorly directed facet for the articulation of the tuberculum of 
the atlantal rib; dorso-medially it articulates with a facet on the 
odontoid-like pleurocentrum; near its ventro-posterior border it is 
perforated by a foramen (vertebrarterialis) ; its ventro-anterior edge 


Parevasaurian Studies.—Part XI. 55 


took part in the articulation to the skull; in the median line the two 
halves of the arch just fail to meet and no spine is developed as is 
primitively the case (Seymouria). The atlantal intercentrum is the 
largest in the entire column; articulating with the odontoid with its 
dorsal surface and laterally with the arch, it forms the ventral half 
of a disc; a posteriorly directed facet for the capitulum of the atlantal 
rib is situated on its postero-lateral edge. In its most primitive 
form such a facet is carried on a distinct process (Seymouria). All the 
cervical intercentra are unpaired. In Procolophon they are paired. 

The pleurocentrum is odontoid-like in shape, but still bears features 
of an embolomerous pleurocentrum; its posterior surface is deeply 
concave and it is only in some cases that there appears to be an 
incipient fusion with the axial centrum ventral to the neural canal; 
its anterior face is strongly convex—the point articulating with the 
concavity in the basioccipital, whereas laterally and ventrally the 
intercalation of the paired arch and the atlantal intercentrum 
complete the articulation with the skull. It is thus apparent that 
the movement of the skull relative to the atlas was mainly rotatory. 

Axis (Pl. XXVIII, Text-figs. 1, 2, and 3).—In the axis the paired 
neural arch is fused with the pleurocentrum to form the reptilian 
type of holospondylous vertebra. The two elements of the neural 
arch are fused and dorsally form an antero-posteriorly elongated 
comb-like spine, which is distinct from that on all the other 
vertebrae; antero-dorsally an upwardly directed prezygapophysial 
facet gives articulation to the atlantal arch; antero-medially at 
the base of the spine there is a paired flange which may be 
termed a zygosphene although there is no corresponding zygantrum; 
postero-dorsally horizontal upwardly facing postzygapophyses give 
articulation to the succeeding vertebra; slightly below the junction 
of arch and centrum there is situated a slightly raised diapophysis 
for the tuberculum of the rib. The centrum is deeply amphicoelous 
and anteriorly is freely articulated with the odontoid which in its 
dorsal part, however, does in some cases show indications of 
incipient fusion; laterally below the diapophysis the centrum is 
hollowed out, ventrally there is a slight ridge; on the antero-ventral 
edge there is no parapophysis for the caprtulum of the rib—this 
facet 1s carried by the axial intercentrum. The axial intercentrum 
is smaller than the atlantal, but otherwise similar. 

In the nature of its neural spine the axis thus possesses a specialised 
character, whereas the deeply amphicoelous nature of its centrum 
without a parapophysis is a primitive character. 


56 Annals of the South African Museum. 


3rd, 4th, and 5th Cervicals.—These three vertebrae are very similar; 
the centra are amphicoelous, laterally excavated, and with a medio- 
ventral ridge; the diapophysis gradually shifts higher up on to the 
arch, and the parapophysis is now situated on the extreme anterior 
edge and gradually shifts higher up on to the centrum; the pre- 
and post-zygapophyses are well-developed and lie in a horizontal 
plane; with each succeeding vertebra the processes bearing the 
zygapophyses extend farther laterally; the centra similarly increase 
in length. The 3rd vertebra differs from the whole series in having 
a pointed pyramidal neural spine; the spines following are massive, 
subquadrate in section, and taller and more slender than the mid- 
dorsal spines but shorter than the first 3 dorsals. The intercentra 
of the 3rd, 4th, and 5th vertebrae are smaller than the two previous 
ones, and the parapophysis has shifted on to the anterior edge of the 
centrum. 

The narrowness across the zygapophyses, the tall spines, and the 
relatively lower centra produce an appearance of tallness in the last 
cervicals in comparison with the dorsals, where the massive zygapo- 
physial and transverse processes cause these vertebrae to appear low 
and squat. 

Dorsals (Pl. XXVIII, Text-fig. 4).—The series of 15 dorsal vertebrae 
present great uniformity in all essential points of structure. Hach 
vertebra consists of an amphicoelous centrum; the anterior and 
posterior faces are, however, not quite similar, viz. the anterior face 
is typically symmetrically concave with the centre as deepest point, 
the posterior face, however, has its upper half concave and its lower 
half convex; the sides of the centrum are excavated and ventro- 
medially there is no ridge or keel; the neural arch is wide, low, and 
massive, and carries a low stout spine; the pre- and post- 
zygaphophyses are horizontal and are carried on widely separated 
processes ; the transverse process for the rib is composed of the fused 
diapophysis and parapophysis, with which is also fused the process 
bearing the prezygapophysis; the articulatory surface for the rib is 
better developed in its upper portion, medially it is not over-functional, 
whereas the lower portion is functional but less developed than the 
upper portion; the transverse process, and in particular its upper 
portion, extends farther laterally than the zygapophyses. Medially 
on the posterior surface of the arch there is a ridge continuous with 
the postzygapophysial process which appears to be an undeveloped 
hyposphene facing an equally undeveloped hypantrum on the 
succeeding vertebra. ‘The differences in the series are of degree; the 


Pareiasaurian Studies.—Part XI. ay 


Trxt-FIc. 4.—Dolichopareia angusta. S.A.M., No. 6238. x 75- 


a=anterior view of a mid-dorsal vertebra. 
b=posterior bes 


99 99 


H. 1=undeveloped hypantrum. 


c=lateral (right) view of a mid-dorsal vertebra. 
d=dorsal 2” 9 29 2? 
H. 2=undeveloped hyposphene. 


yy 


58 Annals of the South African Museum. 


anterior 3 spines are taller than the middle 2-3, and from there to the 
last dorsal the spines again gradually increase in height; similarly 
the transverse process gradually shifts higher up on to the arch in 
each succeeding vertebra, and also increases in length up to the 9th 
dorsal and then decreases; the upper portion also progressively shifts 
in posterior direction, so that whereas in the first dorsals the process 
stands vertical, in later ones it stands obliquely. 

Small intercentra are present throughout the series except in the 
first 5 dorsals, where they appear to be variable. 

As the vertebra immediately anterior to the sacrum bears a rib- 
facet, there can be no question as to the non-existence of lumbar 
vertebrae. 

Sacrals—The number of sacrals is 4; although in the majority of 
cases they are all coalesced, this in not invariably the case. In this 
specimen (S.A.M., No. 9168) of Bradysaurus seeley: there are 4 vertebrae 
with coalesced centra; the zygapophyses are still preserved, but 
could not have allowed any intersacral movement; each bears a rib 
which is firmly fused mainly to the centrum but also in part to the arch; 
the first is very massive with an extended surface of attachment to the 
ilium—its shape is best seen from the illustrations (Pl. X XVIII and text- 
figs. in the paper on the Hind Limb); the second is much less massive 
and is a fairly straight rod-like bone with expanded ends; the third 
is of the same type but more slender; the fourth is still more slender 
and did not reach the ilum, but was ligamentously attached to it. 
The sacrals are much lower than the last dorsal; this is due to a 
decrease in the height of the centrum more than to that of the spines; 
the spines are less massive and consequently appear taller—the first 
two are still subquadrate in section whereas the last two have become 
triangular. The width across the postzygapophyses abruptly becomes 
much less than across those of the last dorsal. 

Pygals.—There are 3 postsacral vertebrae with ribs but without 
chevrons. The ribs are fairly long and slender and are fused low 
down on the centrum, apparently only with the parapophyses. The 
centra are shorter and narrower than in the preceding sacrals, and the 
neural spines relatively longer and more slender and are laterally 
compressed. In all the postsacral vertebrae there is no indication 
of an hyposphene and hypantrum, and all the centra are deeply 
amphicoelous. | 

Caudals.—There are 36 caudals, of which the first 31 bear paired — 
chevrons. The caudals differ from the pygals only in matters of 
degree, viz. in posterior direction they become progressively shorter, 


Pareiasaurian Studies.—Part XI. 59 


lower, and narrower as regards centra and neural spines, and the fused 
ribs eventually become wart-like protuberances. The only delimiting 
feature is the presence of chevrons; these commence with the 4th 
postsacral; they are paired rod-like elements that articulate with a 
swollen proximal end to a facet formed on the ventro-posterior edge 
of the centrum; proximally they enclose a space (haemal canal) 
through which the vascular elements of the tail pass. The zyga- 
pophyses are well developed in the anterior part of the series; they 
are less rigid than in the dorsals; progressively, however, they become 
less and less developed until finally they are absent from the 26th 
postsacral onwards; the last 13 vertebrae articulate only by their 
centra; the last 4 are reduced to centra by the loss of the spine; in 
the last 15 postsacral there are no indications of the transverse 
protuberances to which in the anterior part of the series the proximal 
ends of the caudal ribs are fused. 

In the one specimen (8.A.M., No. 9168) with a complete vertebral 
column the presacral part is somewhat longer than the tail. The 
proportions are: presacrals 13-9, sacrals 2:5, and postsacrals 10-7. 
It thus appears that the tail is longer and comparatively more slender 
than it was formerly thought to be. 

Robs (Text-fig. 5).—All the vertebrae—from the axis to the sacrum— 
bear a pair of ribs. There are thus 20 pairs of presacral ribs. In 
the 5 cervicals the ribs are fairly short (107 mm. in the 4th), slender, 
and dichocephalous—the tuberculum being separated from the 
capitulum by a deep notch. The atlantal and axial ribs have their 
capitula articulated to a facet on the atlantal and axial intercentrum 
respectively. In the next three cervicals the capitula articulate with 
facets on low parapophysial processes situated on the extreme 
antero-lateral border of the respective centra. From the 6th—20th 
vertebrae the ribs are stout and longer—a web of bone connects the 
tuberculum and capitulum so as to produce a holocephalous rib 
which articulates with the long facet produced by the fused diapophysis 
and parapophysis. From the 6th to the 10th the thoracic ribs 
increase in length and in the width between tuberculum and capitulum. 
From the 11th to the 20th the ribs again shorten and become more 
curved. These last ribs curve directly downward, whereas the anterior 
thoracic ribs are directed somewhat posteriorly so that they curve 
obliquely downwards. The last thoracic ribs have a truncated distal 
end which is developed into the nature of a facet; in one anterior- 
thoracic rib the same condition has been observed, and in this case 
it is articulated to a rod-like element bearing a similar facet. It 


60 Annals of the South African Museum. 
MEASURE 


Bradysaurus 
y Nochelesaurus| Embrithosaurus 


baint. seeleyr. | vanderbylt. strubent. schwarz. 
Length of cervical series . ; 280 278 ? 255 Ss 220 
A , dorsal ae ‘ sim iLO) 1115 1090 a0 1065 
i ,», sacral 4 5 A 297 250 250 236 230 
a », postsacral ,, ; : Me 1070 ae at ie 
Atlas— 
Total height é : : ae a 90 a a 
Height of centrum ; : 70 ey 55 67 65 
Width of centrum : : 75 oe 60 73 75 
Length of centrum ; : 50 S 40 51 38 
Height of arch. ‘ ; ui - 74 89 ie 
Length of arch. : : se = 70 69 
Axis— 
Total height : : : 175 144 140 143 ie 
Height of centrum ; q 80 70 60 eel 75 
Width of centrum : : 90 64 65 67 78 
Length of centrum ; : 50 a5) 45 48 45 
Width over prezygapophyses. 100 66 90 76 75 
Width over diapophyses : 110 81 100 100 90 
Width of spine. : 50 41 50 ae as 
Length of spine . ; ; 80 75 70 
| Height of spine . : : 60 60 65 
ord Cervical— 
Height of centrum ‘ Z 82 70 65 7 75 
Width of centrum t : 73 64 60 69 72 
Length of centrum ; : 45 54 50 50 50 
Width over prezygapophyses. 72 80 66 80 70 
Width over diapophyses 5 $7 3 100 105 120 
1st Dorsal— 
Total height : : : 220 193 163 oi 210 
Height of centrum : 5 87 84 75 he 90 
Width of centrum ‘ ‘ 81 80 70 ce 82 
Length of centrum : : 60 61 66 fa 55 
Width over prezygapophyses. 90 7) 80 ee 90 
Width over Giopeu nse: : 145 131 120 ze 150 
Width of spine. } e 40 os ae 62 
Length of spine . : ‘ Pe 32 2 oH, 50 
Height of spine. ‘ : + 51 “ft be 70 
6th Dorsal— 
Total height : : : 205 182 180 220 194 
Height of centrum : ‘ 85 81 65 85 80 
Width of centrum : : 87 80 70 93 82 
Length of centrum : : 60 72 72 67 81 
Width over prezygapophyses. 200 181 148 ? 230 164 
Width over diapophyses : 211 222 153 ? 243 195 
Width of spine. ‘ : 50 55 ws A5 50 
Length of spine . ; ; AT 65 ae 55 55 
Height of spine. : : 63 60 Ae 83 65 
1st Sacral— 
Total height 4 : ; 180 176 187 251 235 
Height of centrum : : 65 55 61 90 70 
Width of centrum t : 69 74 98 97 100 
Length of centrum : : al 76 63 67 80 
Width over prezygapophyses. 193 191 175 207 195 
Width over diapophyses : 137 132 138 183 150 
Width of spine. . ; 66 70 he 60 65 
Length of spine . : : 54 60 - 33 65 
Height of spine. : : 73 75 Ey: 84 70 
6th Postsacral— 
Total height : t : ae 136 
Height of centrum : : 56 45 
Width of centrum ; ; 60 40 
Length of centrum Z ; 37 39 
Width over prezygapophyses. 67 55 
Width over diapophyses ; 93 76 
Width of spine. g : ae 38 
Length of spine . : : = 34 


Height of spine . ; : ae 53 


g 


Pareiasaurian Studies.—Part XI. 61 
MENTS. 
Pareiasaurus LECT ISAS Dolichopareia| Brachypareia | Propappus Anthodon 
serridens. nomen | ieee angusta. rogerst. parvus. serrarwus 
201 160 170 
i 1020 980 a Se si — 
203 150 WAT be af, Sf 147 
105 fe 
65 45 
70 55 
40 35 
68 
95 
57 ae 
43 40 
36 33 
63 49 
7d se 
68 A 
AT 45 
45 43 
70 61 
76 
173 19) 125 
65 61 67 
ie! 69 80 
45 51 oT 
87 74 107 
166 ne 109 
80 es 
AT a: 
55 48 
163 130 219 134 113 112 
61 82 S13) 50 46 
65 64 100 60 S13) 43 
52 64 67 AT 55 Al 
128 155 200 121 105 107 
154 154 223 136 110 109 
42 47 46 42 28 
56 61 45 35 32 
38 84 43 30 33 
163 138 106 me 95 
alk 54 53 65 40 
86 67 63 55 39 
60 50 AQ 50 38 
175 159 158 LEON? 113 
120 135 134 90 83 
69 51 50 32 
58 53 55 33 
50 37 30 29 


VOle xox. PART 1 10 


G2 Annals of the South African Museum. 


would thus appear that some of the ribs were segmented. Un- 
fortunately I have no complete series and cannot determine the full 
nature of this segmentation of the ribs. On the posterior edge of the 
thoracic ribs an irregular flange is developed to house the insertion of 
muscles (external oblique abdominal, sacro-spinal, and the inter- 
costals) ; on the anterior proximal edge a flange received the latissimus 
dorsi and serratus muscles. 


TExtT-Fic. 5.—Pareiasaur ribs. x t. 


I =atlantal rib. 

II =5th cervical rib. 

III =I1st dorsal rib. 

IV =mid-dorsal rib. 

V =last but one dorsal rib. 
c =capitulum. 

t  =tuberculum. 


As has already been mentioned, there are 4 sacral ribs; the first 
is very strong, having a long surface of anchylosis on the centrum 
and has a still greater distal expansion which clasps the anterior 
inner surface of the iliac blade; the succeeding three progressively 
decrease In size and the 4th appears to be only ligamentously attached 
to the iliac blade. 

The anterior postsacrals have fairly long and slender ribs fused to 
the parapophysis. These progressively decrease until eventually they 
are only wart-like protuberances and are finally absent in the last 
15 vertebrae. 

Systematic.—As is the case in the other parts of the skeleton, the 
vertebral column of the various genera and species show only differ- 
ences of degree and not of kind. As a further complication there is 


Pareiasaurian Studies.—Part XI. 63 


the fact that within a single column there are fairly similar differences 
between vertebrae in different parts of the series. Unfortunately 
the column is insufficiently preserved in many species, with the 
result that some characters which may have been used in a classi- 
ficatory scheme are only known in some species. The main points 
of difference are of size, nature of neural spine, proportions of the 
constituent parts, and some rather indefinite variations in shape. 

In our paper on the Hind Limb certain alleged differences in the 
sacrum were stressed. An examination of additional material seems 
to indicate that these points are invalid. The Pareiasaur sacrum 
consists of 4 vertebrae with their ribs, and the differences formerly 
mentioned are preferably to be ascribed to individual and age 
variations rather than to specific differences. In the mature 
Pareiasaur the 4 centra are coalesced and the first sacral rib is the 
strongest; in immature animals 2-3 centra only may be coalesced. 
In the genus Bradysaurus these variations are very well shown. 


Genus BRADYSAURUS. 
(Pl. XXVIII.) 


There are 5 cervicals, 15 dorsals, 4 sacrals, 3 pygals, and 36 caudals. 
The dorsals are large and massive, with strong, though not very high, 
transverse processes. The neural spines are stout and low, with 
somewhat expanded tops, and are roughly subquadrate in section, 
but the four sides of the spine are rounded and not separated by 
very sharp edges. 


Genus NOCHELESAURUS. 
(Text-fig. 6.) 


The dorsals differ from those of Bradysaurus in their absolute and 
proportional greater width across the transverse processes and the 
zygapophyses. The sides of the spines appear to be less rounded and 
taller than in Bradysaurus. 


Genus EMBRITHOSAURUS. 


Very similar to Bradysaurus, except that the last cervicals and first 
dorsals appear to be taller; the vertebrae are, moreover, slightly 
smaller and the width across the zygapophyses is proportionally less. 


64 Annals of the South African Museum. 


Genus DoLICHOPAREIA. 


Very massive neural arch, but with the neural spine taller than in 
Bradysaurus; the two anterior and two posterior faces of the spine 
are flat and are separated by sharp and prominent edges. The 
articulatory face on the transverse process is very long, and the 
width across them is proportionally greater than in Bradysaurus, 
notwithstanding the greater height of the neural spine. 


TExtT-FIG. 6.—Nochelesaurus strubeni. S.A.M., No. 5590. Anterior view 
of a mid-dorsal vertebra. x 


Ble 


Genus BRACHYPAREIA. 
The dorsals are smaller (3) and also relatively less massive than in 
Bradysaurus; centra are deeply amphicoelous; the spine is massive 


and very low, with fairly flat faces separated by ridges. 


Genus PROPAPPUS. 


There is a decided difference in the vertebrae of the two species. 
Those of omocratus are very similar to those of Brachypareia, but the 
spines are even lower, whereas in parvus the spines are much more 
slender and consequently do not appear so low as they really are. 
Moreover, the dorsals of parvus are considerably smaller (shghtly 
more than 4 of Bradysaurus), and the transverse processes are situated 
high up on the arch.and not partially on the centra, and carry a short 
articulatory face. The centra are very deeply amphicoelous and 
in some vertebrae actually notochordal. 


Pareiasaurian Studies.—Part XI. 65 


Genus PAREIASUCHUS. 


There are 5 cervicals, but only 14 dorsals, 4 sacrals, and an unknown 
number of caudals. The dorsals are smaller than in Bradysaurus ($), 
but larger than in Propappus parvus. The spines are tall and not very 
massive; the articulatory face on the transverse process is fairly short. 


Genus PAREIASAURUS. 


The vertebrae are large but not so massive as in Bradysaurus. The 
neural spines are massive and fairly low and are subquadrate in 
section. The articulatory face on the transverse process is fairly 
short. In the Stellenbosch specimen the centra of the sacrals are not 
so much constricted as Watson found in the type material in the 
British Museum. 


Genus ANTHODON. 
(Text-fig. 7.) 


In the specimen from Dunedin (S.A.M., No. 10074) there is preserved 
a dorsal vertebra, sacrals, and caudals. The dorsal is small (slightly 


TEext-FIG. 7.—Anthodon serrarius. S.A.M., No. 10074. x 4. 


a =anterior view. 
b=posterior view. 
c=lateral (left) view. 


greater than 4 of Bradysaurus), comparatively light; the spine is low 
and fairly slender; the transverse process is situated high up on the 
arch and the articulatory surface is short. On the whole, the vertebra 
is very similar to those of Propappus parvus. The centrum is noto- 
chordal with an hour-glass shaped perforation. 

Short Discussion.—lf we are correct in stating that the Pareiasaurs 
possessed 5 cervicals this fact would indicate a definite advance from 
the primitive embolomerous condition where the neck at most 
consisted of 2-3 vertebrae; and they would occupy an intermediate 


a 


66 Annals of the South African Museum. 


position within the assemblage of Cotylosaurs where the number of 
cervicals appears to vary from 2-9; they also would definitely be less 
specialised than the Pelycosaurs and the Mammal-like Therapsids 
where the mammalian number—7—1s fairly constant. The atlantal 
arch shows a definite advance over the Stegocephalian and Seymourian 
condition in that the development of an atlantal spine does not take 
place, and in this agrees with higher reptiles. The apparently unfused 
odontoid is also fairly primitive, whereas the axis shows most of the 
typical modifications obtaining in all reptiles higher than Seymouria. 
The slender dichocephalous cervical ribs, which, in the two first 
segments, articulate with their capitula to a parapophysial facet on 
the intercentrum, are primitive. In Seymouria alone are these more 
primitive, being carried on definite processes. 

It appears that the original number of presacrals in Amniote 
vertebrates was over 20, probably between 23-25. If this generalisa- 
tion holds good, the Pareiasaurs have undergone a reduction in this 
series which does not appear to be paralleled in most other Cotylosaurs, 
Pelycosaurs, and Therapsids. 

The sacrum, consisting of 4 vertebrae with centra mostly coalesced, 
is more specialised than the primitive condition in which there was 
only a single element, and also than that of most reptiles in which 
there are only 2; the Deinocephalia and Anomodontia, however, are 
often parallel with 3-7 sacrals. 

The number of postsacrals—36—is large, though shorter than the 
presacral series, and is greater than in some Cotylosaurs, e.g. Capto- 
rhinus and Labidosaurus, indicating a greater functioning of the tail, 
but less than in others, e.g. Limnoscelis. 


] | 

Hii 
ae 
nl 
aa! 
ie hl 
al 
Hil 

it 

HH | 

HW Hy 

ii 

iH 

iH] 

We] 

Wt 

| 

iit 
ai) 
HH) 

mh) 

Wi 

eel 

i! i 

1H 
a 
WH 

} iit 


a 
eta 


ae ; Sa ut Bat a "s pe std se : te a 
Ps 0 i Aes a ti eae ie Le Park Kine Bat “Ed ae / 
' . " vt 7 oe 
» 4 on " < . » . » a‘ “di 
Boe Ss AGchinet ele a fy Pua 
reiph SIS ay ee: Riise ates basis | 
= of ME my . 
: iy ea brs Tm, peel 
i q st AD Snes 
; PAS PS “ wren | 
| 
o ‘ - * 
: : : bs h Pr Sh eid + 


EXPLANATION or PLATE. 


iy! ‘ « 7 rat * x 

Bradysaurus seeleyi. Ss. A. M.,, No. 9168. — ie be ¥ 

The restoration of the atlas is based on 5 other speci 
element i is preserved. | 


a = Lateral (left) view of the entire column. No atteinee 
to indicate the spinal curvatures. Crosp-sections oe 
spines are given. . 

b= Posterior view of the 2nd verkabita takisy’ eDaSA Te ee 

aa ” » Oth ass (last cervical). 2 

a ” ” ” 9th _ oe (4th dorsal). j 
Ee Albus An ae oe) 15th Pr (10th dorsal). 

=a ” a “ ” 20th 3 i , (15th dorsal). 

et, eels nates 7 UIC au, 1 | Set oaual yo 

Wesetay ee I! Sia ee CE cauplad ee eee oe 

my Se gy ash oe OB istidendad)y, 45 

SANS gs 5 jy) 49th J uge*) (29th caudal). 


Sa RS a 


bette ER Regie + 


ae 


“a 


ON Ee ee so 
. ad 7 


Pires 
oak 


WO o 
Se ANN 
“0 


ere 
Aerie 


ae 


agas 


ewe 


Ey 


j 
ay) 


fi y 5 ® 

/ 4 Gy | é 

Oy | AF Lg (ly 
‘ re re 
M4f7 " 
\ " } i) 
I 

{ h ARG 3 ag 


(ol) 


2. Fossil Plants from Fort Grey near East London.—By Prof. 
R. 8. Apamson, M.A., Botanical Department, University of 
Cape Town. 


(With 27 Text-figures.) 


Introduction.—The flora at present existing in South Africa is one 
not only of great diversity and interest but has also been the subject 
of some investigation and speculation in regard to possible origins 
and relationships. Our knowledge of its past history is at present 
rather fragmentary. 

The fossil plants of the Karoo rocks and of the Cretaceous rocks have 
been investigated to a considerable extent, but the Tertiary period 
still represents a largely unknown epoch in South Africa. Fossili- 
ferous rocks of Tertiary or later age are by no means abundant, and 
such fossils as do occur are poor. This renders the investigation of 
any favourable examples a most important link in the building of 
our knowledge. Of the more recent deposits in this country the 
lignite beds at Knysna formed the subject of a preliminary investi- 
gation by Phillips (8. Afr. J. Sci., xxiv, p. 188, 1927). He recorded 
the occurrence of Widdringtonia, Podocarpus, and Curtisia, all of 
which are living at present at Knysna. Some fossil wood described 
by Warren (Ann. Nat. Mus., ii, p. 345, 1912) as Hugenia cordata is 
probably of Tertiary age. 

Of the fossils described in this paper no descriptions appear to 
exist. A brief and rather popular account of the rocks at Fort Grey 
was given in 1868 by Piers (S. Afr. Magaz., Apr. 1868). He made, 
however, no investigation and, indeed, treats them rather scurvily, 
as they are so much less good as specimens than those obtainable 
from the older rocks. 

In view of the fact that several of the fossils here are conspicuous 
objects it is a little difficult to understand how no collector so far has 
given any descriptions of them. 

The present investigation was suggested by Dr. A. W. Rogers, 
F.R.S., lately Director of the Geological Survey of South Africa. 
It was undertaken in the first place with a view to determining the 


68 Annals of the South African Museum. 


probable stratigraphical position of the rocks, which form an isolated 
superficial deposit of quite local occurrence. 

Acknowledgments.—Part of the investigation has been carried out 
with the aid of a grant from the British Association for the Advance- 
ment of Science. 

I take this opportunity of returning thanks to Dr. Rogers for 
directing my attention to these fossils in the first case, for supplying 
me with material he had collected, and for arranging for the sectioning 
of a number of specimens. I have also to thank Dr. Rogers for his 
continued interest throughout the work. 

I am also indebted to the Director of the South African Museum 
for permitting me to examine a number of specimens in their collection. 
I have to thank Dr. J. V. L. Rennie, who spent some time with me 
in the field, for his assistance both in the collection of the material 
and for suggestions, geological and other. Finally, I have to thank 
Professor A. Young of the University of Cape Town for carrying out 
the sectioning of rock specimens. 

Locality.—The fossil-containing rocks occur on the summits of two 
or three koppies on the south bank of the Buffalo River near Fort 
Grey, about 12 miles from East London. The portion where most 
of the material examined occurs is on Farm 36, known as Spring- 
fontein. I should lke here to express my thanks to Mr. L. W. 
Gower, owner of the farm, for the readiness with which he granted 
free access to his land and for his assistance in various ways on 
visits made to the farm. 

The koppies where the rocks occur form part of the dissected 
peneplain of this region. The fossiliferous rocks are hard and 
silicious and overlie the Karoo Dolerites which form most of the 
surface in the region. 

The main koppie rises to a height of about 225 m. (750 feet) above 
sea-level. The top is a flattish area sloping gently to the south, in 
length about 305 m. (340 yards) and 81 m. (90 yards) in width at its 
widest part. Except on the south, the ground falls away abruptly in 
a low cliff at the sides. 

Rocks.—These superficial silicious rocks vary both in thickness and 
in character. On this koppie the rocks thin out from north to south. 
At the southern end is a small quarry, at one time used for road 
metal. At this point the rocks form a hard bed about 1-2 m. (4 feet) 
in thickness, which, directly overlies very much weathered dolerite. 
At the northern end the exact thickness could not be determined 
with accuracy, as the base of these rocks is nowhere exposed but 


Fossil Plants from Fort Grey near East London. 69 


covered by a talus deposit. On the cliff at the north end a thickness 
of 7-5 m. (25 feet) is exposed. The summit of the koppie is about 
3-5 m. (12 feet) above the edge of the cliff. Descending, true dolerite 
soil was first struck about 15 m. (50 feet) lower down than the base 
of the exposed rocks. 

These silicious rocks themselves are of more than one kind, a 
general threefold division can be recognised; the lowest rocks seen 
are hard and claylike, blueish when fresh but weathering to a rusty 
brown or red. Above these is a hard, often flintlike rock with no 
regular joints and a conchoidal fracture. This rock is grey or purple 
when fresh, but becomes white with partial weathering. The upper- 
most part is a brown or red rock which varies from a hard compact 
chertlike rock to a granular quartzite. In some parts it looks like 
a sand cemented by amorphous silica. Local bands occur which are 
almost wholly amorphous silica. On the koppie these uppermost 
beds overlap the others at the southern limit and occur directly above 
the basal dolerite. These main divisions can be subdivided, though 
the subdivisions are more local in character. 

Other Localities— The above description is based on the main 
koppie on Farm 36. A similar deposit, but of less thickness and 
containing much fewer plant remains, caps the ridge forming the 
east boundary of the farm. In this the hard flinty bed seems wanting. 
A further small deposit occurs as a cap to a rise about ? mile to the 
south-east, but this has yielded no fossils. 

A general investigation has been made of the surrounding country 
for other deposits of this kind, but so far without result. Both sides 
of the Buffalo River were examined and the study continued in a 
general way southwards to the coast and west to the Keiskama 
River. The only evidence for other deposits in the neighbourhood 
was the presence of some loose blocks of quartzite in a field near 
Need’s Camp, but the origin of these was not found. 

Materval.—The locality was visited in 1930 and again in 1932, and 
a quantity of material was collected. The bulk of this has been 
deposited in the South African Museum. In addition some specimens 
were obtained from Dr. Rogers, and the material in the Museum was 
examined. 

The material consists of fragmentary plant remains which are 
siligified. The original material has been completely replaced in some, 
but in others a powdery condition is found which is very fragile and 
difficult to work with. In no case is there any carbonaceous material 
or other substance that could be separated from the matrix. 


70 Annals of the South African Museum. 


Fossils occur in all the beds, but in the lowest parts are structureless 
impressions. The hard flinty rock has provided the majority of the 
workable material. Especially in the uppermost layers plant remains 
are often in cavities that suggest shrinkage having occurred. 

Descriptions.—In the following account the remains are grouped 
in accordance with their nature; their distribution in the rocks is 
recorded, but is discussed later. 


GYMNOSPERMS. 


Wood.—The most prominent fossils are woody axes which are not 
infrequent. These are of various sizes, varying from small twigs to 
portions suggesting pieces of large trees. 

The woody axes occur in all the beds of rock, and fragments are 
often found weathered out from the rock in the talus flanking the 
koppie. Some of these separate specimens have been carried down 
some distance beyond the limits of the rocks from which they were 
derived. In size there is considerable variation, from twigs 0-75- 
2-5 cm. (4-1 inch) in diameter up to logs 40 cm. (16 inches) or more. 
Among the largest specimens measured was a weathered-out portion 
which consists of about half the original axis and which has a 
diameter of over22cm. Another much decayed and weathered portion 
had a length of 4 my 18 cm. and a diameter of 54 cm. In the rock 
these axes occur in all the beds. Some are erect or ascending, others 
horizontal. The former give all the impression of being preserved 
wm situ, an impression which is further strengthened by the fact that 
in some cases an axis can be traced through more than one bed of 
rock. In the quarry at the south end of the koppie these axes are 
continuous down into very much weathered dolerite. They seem 
especially abundant at the top of the lowest beds. The examples 
that are apparently in situ are generally not very large; the largest 
erect stem was a portion 80 cm. (32 inches) in length with a diameter of 
18-75 cm. (74 inches). Much longer portions of smaller diameter occur. 

The external appearance shows a good deal of variation; some are 
quite smooth, others ridged, or fluted in various ways. Little weight 
can be put on these external features, as practically none of the 
specimens are complete. The wood only is preserved, the original 
outer tissues are wanting. The wood itself in many of them has been 
eroded before fossilisation took place. 

Some of these axes show branching. The branches occur singly, 
not in whorls. 


Fossil Plants from Fort Grey near East London. 71 


Structure.—As seen in hand specimens the structure in all the axes 
is very uniform and certainly that of a Gymnosperm. The wood is 
made up of uniform-sized tracheids, and is traversed by numerous 
narrow and shallow medullary rays. In the majority annual rings 
are not distinctly present and sometimes quite absent. 

A few specimens exhibited an appearance of broad rays at intervals 
between the narrow ones, but more detailed study showed that this 
was the result of radial splits or cracks that had become filled with 
silica. 

The degree of preservation of the internal structure shows a good 
deal of variation; in a few, much of the structure is preserved, 
though generally the finer details, such as pitting on the walls, are 
faint and ghostlike. 

Some thirty specimens have been studied by means of sections or 
transfer films. To avoid repetition the general features common to 
all the specimens will be given first, and the details that separate 
the different kinds of wood later. 

The wood is made up of very uniform tracheids, between which 
are numerous medullary rays which are one cell in width. Among 
the tracheids are vertical rows of cells filled with a dark material. 
These cells are rarely well preserved, but appear to have been shorter 
than the tracheids and represent parenchyma. 

In the larger specimens annual rings are not at all distinct and 
sometimes quite absent. Of 26 transverse sections, 15 showed no 
signs of annual rings, 7 had very indistinct rings, more obvious in the 
hand specimen than in the section, 2 had rings that were separated 
from one another by a single line of smaller tracheids, 1 showed 
rings of this kind in the outer parts but not near the centre. One 
specimen had been so much compressed that no conclusions could 
be drawn. 

These differences in the formation or otherwise of annual rings 
could not be definitely correlated with differences in other structural 
features. 

While annual rings are not distinct, many of the specimens, and 
especially the larger ones, show a pronounced concentric zonation, 
which is due to compression. Zones round the axis occur in which 
the tracheids have become squashed and more or less displaced. 
These compression zones are readily distinguished from annual rings 
in a section owing to the distribution of the medullary rays. 

The numerous medullary rays are composed entirely of parenchyma; 
in depth they are most commonly 2-6 cells, though occasionally 


712 Annals of the South African Museum. 


deeper rays, 8-12 cells or even 20 cells, occur. An appearance of 
even deeper rays may be given by the close approximation of numerous 
small rays (fig. 1). 

The tracheids forming the wood show differences in size and thick- 

ness of the walls in different specimens. These differences seem 

largely due to the state of preservation and to 
ee the condition of the specimen when preservation 
(ie Deere took place. In some, pits are easily visible, 
LE gE? in others they are very faint or absent. There 

are bordered pits on their radial walls. The 
Fic. 1.—Long.tang.sect. bordered pits are circular, and occur in a single 

wood showing medul- : : 

lary rays. x55. row, or in two rows, when the pits are oppo- 

site to one another. In some cases one tracheid 
may show bordered pits, both paired and single, and in such cases 
occasional alternating pits are found. The bordered pits may be 
separate or in contact. In the latter case they may be somewhat 
flattened along the line of contact. Polygonal pits do not occur (fig. 2). 

Thickened rims, rims of Sanio, between the pits are generally 
present and sometimes very distinct. In some they seem absent, 
but as these are the least well preserved the negative evidence is of 
little value. The pits on the tracheids occur in groups towards the 
ends of the cells and in the neighbourhood of medullary rays. 

No resin canals are present either in 
the wood or in the medullary rays. 

In most of the specimens wood alone 
is preserved, but in a few the central 
parts are preserved: both stems and 
roots occur which show no noticeable 
differences apart from the primary 
structures. 

In stems the central pith is of small 
size and parenchymatous with cells 
rounded in cross-section and not much 
elongated longitudinally. The primary 
xylem projects into the pith. No 
specimen shows any detail in the primary wood. | 

The roots have a solid core. The primary xylem appears to be 
4-arch, but this may be due to compression and poor preservation. 
Small rootlets associated with the larger axes are 2-arch. 


Fig. 2.—Long. rad. sect. wood 
showing pits. x 80. 


Both in stems and roots the outer tissues are very seldom present 
and are absent in all the larger specimens. Even where bark is 


Fossil Plants from Fort Grey near East London. 73 


found very little detail can be made out. The bark seems to have 
been thin and smooth. Cork occurred on the surface. There is no 
evidence for the presence of resin canals. 

It is possible to distinguish three types of wood. 

The first type, which is much the commonest, has parenchyma 
scattered among the tracheids without regular arrangement. The 
rays in transverse section are narrower than the tracheids. Annual 
rings are never distinct. The tracheids have one or two rows of 
bordered pits separated by rims of Sanio, and connect to the rays 
by one or two pits. These field pits are bordered and circular in 
outline: the central aperture was probably, but not certainly, 
elliptical (fig. 3). 

A few specimens only differ from this type by possessing rays of 
much larger cells, as wide as or wider than the 
tracheids. 

The second type of wood 
is less common. The rays 
are narrow. Not very dis- 
tinct annual rings are pres- 
ent. The ray cells have 
three or four rather small Fic. 4.—Sect. similar 
Lee field pits (fig. 4). The wood eee Pipe 

showing field pits, Contains less parenchyma, from a photograph.) 


x125. (Traced from which is often in tangential 
a photograph.) 


bands running between the rays. 

The third type has only been found in small specimens, not more 
than 5 cm. (2 inches) in diameter. The most obvious character is 
the presence of well-marked annual rings. These are narrow and 
separated from one another by zones of very small tracheids forming 
the autumn wood. Throughout the tracheids are smaller than in 
the other types, and the appearance is given of a harder more compact 
wood. In detailed structure the differences are less; a small amount 
of parenchyma is scattered through the wood, the uniseriate rays 
are 2-4 cells deep or occasionally 6-8 cells, bordered pits are circular 
in One row, contiguous but not flattened. Rims of Sanio are in- 
distinct. The field pits are 1-2 and circular. The pith in the 
centre was of cells with thickened walls. 

Branching occurred freely in these stems; several specimens have 
an irregular structure due to the formation of branches as large as 
the main axis. 

Comparisons.—Of these fossil woods the first type seems to show 


74 Annals of the South African Museum. 


a close agreement with Podocarpus. There are some points of dis- 
agreement: in the fossil the diameter of the tracheids is larger than 
in recent Podocarpus wood, and the bordered pits appear to have a 
circular aperture while the recent wood has an oblique elliptical one. 
Both these differences may be explained by lack of perfect pre- 
servation. 

The second type, characterised by less parenchyma and grouped 
field pits, is more like Widdringtonia than Podocarpus. Both these 
comparisons are supported by other structures, especially leaves and 
young shoots that occur in the rocks. — 


STRUCTURES ATTRIBUTABLE TO Podocarpus. 


Leaves.—In the uppermost portion there occur impressions of 
lanceolate parallel-sided leaves which have a definite central vein 
but no lateral or branch veins. None of the impressions are complete, 
though some show signs of a narrowing at the ends. The largest 
is 2-8 cm. long and 9 mm. wide. These impressions have no structure 
preserved. 

Another specimen is a silicified fragment of a linear lanceolate leaf. 
A part only is exposed, 4 mm. long and 1-3 mm. wide. The fragment 
tapers to one end, which is apparently the base. At the broader part 
there is a faint ridge in the centre which is about a third the width. 
Transfer films show the surface features. The 
epidermal cells are rectangular, rather elongated 
along the midrib but not at the sides, where there 
are rather large stomata arranged in straight 
rows. The guard cells are sunk below the surface 
(fig. 5). 

This leaf is certainly that of a Gymnosperm, 
and, except for the very small size, agrees in its 
characters with Podocarpus. 

In several sections are fragments of small leaves 
Fic. 5.—Epidermis which are not well preserved. These have a 

aes leat. yalisade layer occupying about half the thickness. 

Vascular bundles are generally not visible, but 
some show a thickening in the middle region. One or two show 
indications of a hypodermal layer on the upper side. 

Seed.—In the upper layers two specimens have been found which 
may be seeds. The more perfect has been split open. It is rather 
flattened, about 8 mm. long and 4-5 mm. across. It is pointed at 


Fossil Plants from Fort Grey near East London. 75 


one end, but not symmetrically so; one side is straight, the other 
distinctly curved. The widest part is near the base. There is a 
smooth central mass surrounded by an outer wall. The central mass 
at its widened base has a sharp conical projection, 0-5 mm. long, 
opposite the angle of the bulging side. This structure immediately 
suggests a comparison with the seed of Podocarpus. The central 
mass with its projecting point is the prothallus enclosed by the 
nucellus with a nucellar beak. 

The other specimen, found in proximity, is complete and shows 
the external surface only. Itis 12 by 9 mm., with a rounded outline, 
very slightly pointed. The surface at the edges where flattening has 
occurred is slightly grooved. 


STRUCTURES ATTRIBUTABLE TO Widdringtonia. 


In various parts of the upper beds are portions of shoots bearing 
small leaves in decussate pairs. In the younger parts these leaves 
are closely appressed, but in older eas they are longer and with 
free projecting tips. These cupres- 
soid twigs may be silicified, but are 
more commonly impressions on the 
rock. 

Where the leaves are appressed 
they appear broadly triangular with 
a pointed or rounded tip. The 
spreading leaves are ascending or 
squarrose: the free portion is 


Fic. 6.—Tracing of 
cupressoid twig. Fic. 7.—Tracing of 
x 4. cupressoid twig. x4. 


flattened on the adaxial side. In the majority of cases the leaves 
on a twig are of uniform size, but in a few smaller leaves occur at 


the base (figs. 6 and 7). 


76 Annals of the South African Museum. 


Some of the specimens show internal structure. They are un- 
doubtedly Gymnosperms, though details of wood structure are lacking. 

The leaves in section are plano-convex, rhomboid, or curved: at 
the tip nearly circular. Each is traversed by a single vascular bundle 
which is central or nearer the upper side. In the lower half a large 
resin canal follows the bundle on its abaxial side (fig. 8). 

Palisade tissue is not very regular but is more developed on the 
abaxial surface. The epidermisis thickened, but there is no hypoderm. 

In one specimen traces of stomata of the Gymnosperm type 
occurred on the adaxial face. 


Fic. 8.—T.S. of leaf from cupressoid twig. Fic. 9.—Possible male 
x 80. cone. x40. 


Possible Cones.—In one section in close proximity to leaves of the 
cupressoid type there is a rather vague structure that may be a 
fragment of a male cone. It consists of a short axis bearing three 
structures that widen to the tips: two are lateral, one seems terminal. 
Two of these bear hollow structures attached below the expanded 
tip. The whole structure is about 0-75 mm. in length, and does not 
show good cellular structure (fig. 9). 

In another piece of rock containing cupressoid twigs is a flattened 
body that may be a cone scale. This is about 2 cm. long, and is 
blunt-ended and narrowed to its base. The surface is smooth and 
marked by faint lines that converge towards the blunt apex. The 
specimen shows traces of cell structure on its surface. The epidermis 
is composed of small rectangular cells, except along the lines mentioned, 
where they are elongated. No stomata were visible. 

Gymnosperm Roots.—In some parts of the rock there occur numerous 
roots with characteristic gymnosperm structure. These are from 


Fossil Plants from Fort Grey near East London. Uh 


2-6 mm. in diameter. In transverse section they consist of a central 
mass of tracheidal wood traversed by narrow medullary rays of 
parenchyma. The primary xylem in the centre is diarch. There 
are no resin canals. The outer tissues are present, but are in a poor 
state of preservation (fig. 10). 

All the roots have secondary tissues developed, though in some of the 
smaller ones the amount is not large. In the larger ones no division 
of the wood into annual rings is noticeable. 


Fetnye 


Fig. 10.—T.S. gymnosperm root. x65. 


These roots have not been found in actual connection with any 
larger axis. 
FERNS. 


Remains of ferns occur somewhat locally in the rocks. While not 
generally distributed these remains are abundant in patches, especially 
in the upper parts of the hard beds. 

All the fern remains found are fragmentary and, at present, do not 
allow of certain identification, or even of clear association of different 
portions of the plants. 

Ferns are represented by stems, roots, petioles, fragments of leaves, 
and traces of sporangia and spores. 

Stems.—Stems of ferns of more than one kind have been found. 

Solenostelic Stems.—These are the largest, and, while only found in 
one portion of the rock, are abundant there. 

These stems may be as much as 10 mm. in diameter and are com- 
monly 6-8 mm. They occur generally as short fragments, no piece 
over 2 cm. in length having been found. In section they appear 
elliptical, but this seems due to compression. 

VOln xXx, PART I, Il 


78 - Annals of the South African Museum. 


These stems show typical solenostelic structure, a closed ring stele 
with a central pith region and a relatively wide cortex outside; the 
stele is bounded both outside and inside by phloem, pericycle, and 
endodermis. The xylem is made up of tracheids associated with 
parenchyma. In longitudinal section the tracheids have scalariform 
thickenings. 

The cortex in some specimens is uniform parenchyma, in others it is 
differentiated into zones, the inner and outer having thin-walled cells, 
while the middle region is thick walled. The pith may show the same 
differentiation, though the thick-walled portion extends to the centre. 

At the surface the cells are very thick walled. In the better pre- 
served specimens there are projections from the surface in the form 
of rows of cells or of groups of 2-3 cells. These represent ramenta, 
which were abundant. 


Fic. 11.—T.S. solenostelic fern stem showing branching and departure of 
leaf trace. x7. 


Some portions show nodes. Here a small gap is formed in the 
stele: the leaf trace passes out through the cortex in the form of a 
curved strand. 

Branching of these stems occurred. Branching is apparently 
dichotomous, though one division is distinctly larger than the other. 
Roots arise in numbers from these axes. No gap is formed in the 
stele at the departure of the root trace (fig. 11). 

Protostelic Stems.—These are more common than the solenostelic 
stems described above, and are more generally distributed through 
the upper parts of the rocks, though never so abundant in any part. 

These protostelic stems are smaller in size than the solenostelic 
ones. The largest is 5-6 mm. in cross-section, but 0-75-1:5 mm. is 
the more general size. Though the diameter is less these stems may 
occur as continuous axes of 5-6 cm., following a straight course 


through the rock. 
Two types of protostelic axes can be recognised. 


Fossil Plants from Fort Grey near East London. 79 


First Type.—These are generally larger, 2-3 mm. in diameter as 
an average. They exhibit a wide cortex and a central stele. The 
cortex may be uniformly thickened or have the outer and inner parts 
thick walled and the middle region thinner. The epidermis is smooth, 
with no trace of ramenta. 

The central stele, which may be a third or more of the total thick- 
ness, has the middle occupied by xylem. This is surrounded by a 
narrow zone of thin-walled cells, which represents the phloem and 
pericycle. In no specimen is this zone well preserved. There is a 
definite endodermis between this zone and the cortex. 


Fia. 12.—T-.S. exterior of xylem of protostele Fic. 13.—Sect. similar to 12 with 
showing mesarch protoxylem. x 140. exarch protoxylem. x 140. 


The central xylem is made up of tracheids with scalariform thicken- 
ings interspersed with parenchyma. The tracheids may be wide. 
The protoxylems are mesarch (fig. 12). 

One of the largest specimens found includes a node. At one side 
of the stele is an arc of xylem separated from the main mass by a 
patch of rather small celled parenchyma, the cells of which are thick 
walled. In the sections obtained this parenchyma is enclosed by 
xylem, the arc being attached at the sides to the main mass. At 
the opposite side, ventral side, of the stele are projections from the 
xylem that represent root traces. 

Second Type.—The stems included here are of smaller size, 1-5 mm. 
as the average. In general structure they are very similar to the 
first type, but differ in having the protoxylems exarch (fig. 13). Also 


80 Annals of the South African Museum. 


the tracheids in the xylem are differently arranged: instead of being 
scattered with separating parenchyma they occur in groups with the 
cells flattened at the points of contact. 

Petioles.—These are the commonest fern remains, and in some 
pieces of the rock are abundant. No case has been found where a 
petiole is attached to a stem. The examples noticed of nodes in 
stems have only the leaf traces and not the free portion of the petiole. 

These petioles are cylindrical or flattened, and slightly grooved on 
the upper surface. In diameter they vary from quite small structures 
to 2-2-5 mm. Portions as much as 3 cm. long have been found. 

In structure they are fairly uniform, the stele consisting of a 
single continuous horse-shoe-shaped band with central xylem. This 


& (®) Fie. 16.—Outline 


Fig. 14. — Outline Fic. 15. — Outline of petiole with 
of fern petiole. of fern petiole. divided stele. 
x 9. x 9: x 9. 


band may be simple or the tips may be thickened or bent round in 
hooks. Where hooks occur at the tips these may be bent over to 
be parallel with the main portion or spread out at an angle (figs. 
14, 15). 

Some portions have been found in which the stele is divided up 
and has the form of an are more or less curved, sometimes distinctly 
U-shaped, with two separate strands which themselves may be 
curved. There is no significant difference in size between those with 
a continuous stele and those with the divided stele (fig. 16). 

It is not possible to associate these petioles definitely with the 
stems. The different stems occur associated in the same rock. 
Petioles with divided steles have not been found associated with 
solenostelic stems. 

Roots.—Kspecially in association with the solenostelic stems small 
roots are exceedingly abundant. These roots appear unbranched. 
In section they have a small stele with diarch xylem. The endo- 
dermis consists of cells distinctly elongated in the tangential direction. 
The cortex is thin walled (fig. 17). 

Leaves.—The remains of leaves attributable to ferns are exceedingly 
fragmentary and of little value for purposes of identification. 


Fossil Plants from Fort Grey near East London, 81 


Some portions occur as impressions which represent fragments of 
an apparently compound leaf, and probably ultimate segments. 
These are pinnatifid scraps with forwardly directed rounded lobes. 
The venation is dichotomous without anastomoses. No cell structure 
is preserved. 

In some of the sections in association with the petioles and fern 
roots are a number of scraps of leaves. These are thin, flat structures. 
The epidermis is not distinctly thickened. The mesophyll on the 


Fic. 17.—T.S. fern root. x80. 


upper side has a rather irregular palisade formation. The preserva- 
tion is poor and no vascular bundles with structure are present. 

Some of these leaf scraps show a downward curving of the margin 
of the leaf. 

Sporangia.—No sporangia or any trace of them have yet been 
found in direct connection or immediate association with the leaves. 
All that have been discovered are some poorly preserved specimens 
lying isolated in the rock containing fern remains. No complete 
specimen has been found. Some of the sporangia are circular, with a 
rather massive annulus running round the periphery. The annulus 
extends about two-thirds round. The appearance suggests an 
equatorial annulus (fig. 18). Others are more elongated, with the 
annulus extending half or more longitudinally. These seem to have 
had a vertical annulus (fig. 19). 

Spores.—Some groups of spores showing formation on tetrads occur, 
but they do not show any distinctive features. 

Comparisons.—Owing to the fragmentary nature of these fossils, 
especially of the leaves and sporangia, detailed comparisons with 
living plants are difficult. 


82 Annals of the South African Museum. 


The protostelic stems of the first type, with mesarch protoxylem, 
and the pocket of thickened ground tissue at the node, agree closely 
in size and many details of structure with species of Gleichenia. 
Further support is got from a section of an axis closely associated 
with one of these protosteles which has the xylem in a ring enclosing 
a small central area. This is extremely like a section of a petiole of 
Gleichenia formed near a rapidly growing rhizome tip in which the 


Ss 
y ‘ i 
J 
Fic. 18.— Sporan- 
gium with equa- Fie. 19.— Sporan- 
torial annulus. gium with vertical 
x 130. annulus. x 130. 


stele forms a closed ring. These fossils agree in so many features 
with Gleichenia that generic identity may be accepted. 


The identity of the other fern types must remain unsettled at 
present. 


ANGIOSPERMS. 


Both monocotyledons and dicotyledons are found, though all are 
of small size. Most of the fossil material is in the form of isolated 
scraps which are described if only to give a stimulus to subsequent 
collectors. 


Monocotyledons. 


Stems.—Stems of a small monocotyledon are abundant in local 
areas in the rock. These stems are of two kinds, but as they have 
been found in attachment they are from the same plant and represent 
rhizomes and aerial axes respectively. 

Rhizomes.—These are cylindrical structures as much as 2-3 cm. 
long and 2-5-3-5 mm. in diameter: a few reach 5-5 mm. Most are 
unbranched. They have a characteristic appearance due to furrows 
on the surface which run both longitudinally and horizontally and 
divide the surface into rectangular areas which have the longer axis 
horizontal. In the younger portions scale leaves arise from the 


Fossil Plants from Fort Grey near East London. 83 


raised areas and enclose the terminal part: these are wanting in 
older parts. 

In section these axes show the characteristic rhizome structure, 
with a wide cortex and the bundles in a definitely bounded central 
cylinder which is often less than a third of the total diameter 
(fig. 20). In the central cylinder are numerous bundles which follow 
a rather irregular course. The bundles are close at the periphery 
but more separate and larger in the centre. Hach is concentric, 
amphivasal, with a small phloem. The cells of both phloem and 
xylem are narrow (fig. 21). The cortex is composed of thin-walled 
parenchyma cells with rather numerous small spaces between them. 


' Fre. 20.—Monocotyledon Fic. 21.—Amphivasal bundle 
rhizome. x9. from rhizome. x 200. 


The epidermis is a very distinct layer. In old portions regular layers 
of cork occur on the outside. 

Scale Leaves.—In section these seem structureless. It is probable 
that they were dry, shrivelled scales. Near the base they are com- 
posed of undifferentiated parenchyma through which run a number 
of small parallel vascular bundles. 

Roots.—Roots are produced in large numbers from the rhizomes. 
They are found passing through the cortex in all directions and 
emerge indiscriminately through the raised areas and the furrows. 
The roots are small, rarely 0-5 mm., and are unbranched. They have 
a parenchymatous cortex and a relatively wide central cylinder which 
is medullated and has numerous strands of xylem. 

Aerial Stems.—These are slender and slightly compressed and show 
no trace of division into nodes and internodes and have no appendages. 
They may be as much as 2 by 1 mm. in section, but are generally 
smaller, 1-0-1-25 by 0-5-0-75 mm. Internally the cortex is narrow 
and the central region, with scattered bundles, not sharply separated 


84 Annals of the South African Museum. 


from it. The cortex is traversed by strands of subepidermal fibres 
which are widest outside and extend in almost to the bundle region, 
so that the palisade-like parenchyma of the cortex comes to the 
surface along quite narrow lines (fig. 22). The bundles run parallel; 
each is collateral, not concentric, and is surrounded by a sheath of 
fibres. The xylem is spiral, reticulate or pitted, with the pits 
elongated transversely. : 

General.—These monocotyledons seem to have been plants with 
scapose aerial’stems which grew separately, not in tufts. The struc- 
ture of the bundles excludes the Gramineae and almost certainly the 
Cyperaceae also. In the absence of any flowers more certain identi- 
fication is not possible. 


Fie. 22.—Aerial stem Fic. 23.—T.S. monocotyle- 
of monocotyledon. don leaf. x75. 
x 14. 


Monocotyledon Leaf.—In a section from a part of the rock remote 
from that which yielded the plants just described is a portion of a 
transverse section of a leaf that seems that of a monocotyledon. 

The section is incomplete. It shows a slightly curved outline with 
three vascular bundles. The epidermis is of thick cells, the mesophyll 
of regular parenchyma with few intercellular spaces. The cells are 
polygonal and similar on the upper and lower sides. The bundles 
are embedded. Hach has a sheath, 3-4 cells deep, of small cells. 
The interior is poorly preserved, but towards one side are two large 
rounded cells that look like vessels (fig. 23). 

Dicotyledons.—Scraps of stems and leaves, very numerous roots, 
and fruit structures have been found. The material does not allow 
of definite association of the various parts and is in most cases in- 
sufficient for identification. 

Woody Stems.—These are not abundant and are of small size. 
They occur in the hard middle layers and in the upper beds. The 
best preserved are twigs about 5 mm. in diameter. They have a 
rather thick cortex round the wood, which is characterised by the 


aa 


Fossil Plants from Fort Grey near East London. 85 


presence of large vessels which occur singly or in radially placed pairs. 
The vessels are surrounded by small thick-walled cells. Medullary 
tays, 2-4 cells across, traverse the wood (fig. 24). No longitudinal 
sections are available. 

The cortex had fibres in groups. Fibres also occur in the phloem, 
in which the rays widen out. 

Roots.—Roots of more than one kind have been found, and in some 
parts are exceedingly abundant. None have been found in con- 
nection with a stem, so that identification is not possible. 

Four types of root can be distinguished. 

First Type.—These are much the com- 
monest, indeed they are the commonest 
fossils. They occur in all the layers. In 
the lowest, roots are extremely abundant, 
but are in the form of casts or structure- 
less tracks. In the hard rock, parts consist 
of very little else than these roots. The 
roots run in various directions, but are Fie. 24.—Dicotyledon 
most often vertical and at times are very wood. x 105. 
regularly parallel. 

There is some variation in size; an average diameter is 2-5 mm., 
but some are as much as 5-8 mm., and there are many small ones. 
The smaller ones are often in groups. Branching occurs, but not 
commonly. 

In structure the roots have a small central cylinder surrounded 
by a wide cortex of rather regular parenchyma in regular radial rows. 
The cells of the cortex appear rectangular in cross-section and have 
very small or no spaces between. On the outside there are 5-6 layers 
of smaller cells with thicker walls. In longitudinal view the cortical 
cells are rectangular and about 2-3 times as long as wide. 

The central cylinder is bounded by a definite layer which is not 
distinctly thickened. The centre is filled by the xylem, which is 
made up of rather small cells, very uniform in size. There are no 
large vessels. Pith is only found in the largest roots. The xylem 
is typically 5-6 arch, with the protoxylems at the points of the 
pentagonal or hexagonal column. Small roots and branches have 
4—3 protoxylems. 

The xylem cells have spiral and reticulate thickenings. 

The phloem and pericycle are in most cases torn. They had thin- 
walled cells. 

Many of the larger roots show secondary thickening, but none have 


86 Annals of the South African Museum. 


more than the first stages. The secondary xylem is composed of 
small tracheid-lke cells with regular arrangement. 

Stretching and some disorganisation of the pericycle and endodermis 
occur, but in no case is the cortex cast off, nor is there any sign of cork. 

In many of the larger roots the cortex is penetrated by smaller 
ones of the same kind. Three or four of these penetrating roots 
may be found round the central cylinder of a large root. 

Note on the Cortex of Roots.—The very regular large-celled cortex 
with few intercellular spaces is very characteristic of these roots. 
This cortex is very different from that usually associated with roots 
with a wide cortex in which secondary growth takes place slowly 
and not in much quantity. A much more common arrangement is 
one of cells that are rounded or polygonal in cross-section. 

A comparison of these fossil roots with roots of similar size from 
living plants has led to the view that the form of the cells in the 
cortex of the fossils is not the original one but is brought about by 
alterations in the course of preservation. A careful examination of 
the cortical cells in well-preserved specimens of fossil roots reveals 
small projections on the walls and especially on the tangential walls. 
The projections are lines or ridges running along the walls. Careful 
tracing of these lines exhibits a framework of a cell pattern of 
polygonal or rounded cells with intercellular spaces. It may be 
noted that at times this arrangement is more definite in a photograph 
than in direct examination. 

The formation of the regular rectangular cell appearance from cells. 
with rounded or polygonal outlines would seem to be brought about 
by shrinkage in preservation combined with some disruption. The 
frequent breakdown of the phloem and pericycle is further evidence 
of shrinkage having taken place. To test this some experiments. 
were made on living roots. Roots of Villarsia ovata were used as a 
type of a size similar to the fossils. The cortex in this plant is com- 
posed of radial rows of cylindrical cells, rounded in cross-section. 

Various methods of shrinkage were tried, most of which were un- 
successiul, but one very simple method gave striking results. 
Sections or pieces of root were placed in a solution of gelatine that. 
was allowed to concentrate and set by evaporation at room tem- 
perature. While frequently the contraction of the gelatine caused. 
rupture of the tissues, In some cases an appearance in the cortex 
very strikingly lke that seen in the fossils was obtained. The 
originally rounded cells have been pulled out and appear rectangular 
in section and the spaces have been quite obliterated. 


Fossil Plants from Fort Grey near East London. 87 


Second Type.—These occur in association with the first type of 
root, but are much less numerous. They are generally of smaller 
size, 1-2 mm. in diameter. The cortex is less regular and has not 
the zone of thicker cells on the outside. Correlated with this absence 
of the thicker cells the cortex generally appears compressed or de- 
formed in various ways. The central cylinder is relatively larger, and — 
is bounded by a definite endodermis of rather large cells. Internal 
to the endodermis are two or three layers of cells with thin walls. 
The centre is occupied by a mass of small cells which are apparently 
xylem mixed with parenchyma. There are no distinct rays or 
projection from the xylem. 

Third Type.—A few examples of another and distinctive type of 
root have been found. These are small, 1 mm. or less in diameter. 
The cortex has the cells somewhat thickened. The central cylinder 
is about one-third the total thickness of the root. The greater part 
of the central cylinder is occupied by xylem, which here contains 
large vessels associated with smaller thick-walled cells. The xylem 
seems to be tetrarch. The peripheral portions of the central cylinder 
are not preserved. The endodermis appears to have been thick 
walled. 

Fourth Type.—The roots placed here are slender structures about 
0-5 mm.in diameter. The central cylinder is very small, with triarch 
xylem. The cortex is in three distinct layers; an inner zone of small 
cells, a middle region of large cells with very thin walls, and an outer 
zone of smaller thickened cells. The middle zone is in most cases 
poorly preserved and incomplete. 

Comparisons.—While identification is not possible, some general 
comparisons may be made. The first type of root, which is rather 
thick and has very little secondary growth, seems most like the root 
of a perennial herb. Roots very similar in their general features 
occur in some of the Compositae, which have roots which are little 
branched, with a wide cortex, and with small amounts of secondary 
growth. The xylem is also composed of small cells. 

Leaves.—In the uppermost layers of the rock there are a number 
of scraps of leaves which have an astomosing venation. The preserva- 
tion is poor, and no details of epidermis or of stomata are obtainable. 
The leaf is represented by a powdery substance which disintegrates 
at a touch. None of the fragments discovered has a complete 
outline. There seems to be certainly more than one type of leaf 
represented. Some have a single midrib with pinnate venation, 
others apparently more than one main vein. These latter have 


88 Annals of the South African Museum. 


smaller veins running across between the main ones and smaller 
branches forming a reticulation. 

Fruits.—In the uppermost layers are found isolated bodies that 
are fruits and probably fruits of dicotyledons. Most occur as casts 
or impressions without cell structure. 

“ Four-chambered Fruits.’—Two specimens have been found of 
bodies which are almost spherical and smooth externally. These are 
2-4 mm. in diameter. Internally they are divided into four equal 
chambers. The outer wall is $ mm. in thickness; the partitions, 
which run at right angles to one another, are thinner; they are 
slightly thicker at the centre. In one specimen the chambers are 
empty ; in the other, which is silicified, there are indications of con- 
tents. Each chamber held one body, probably a seed (fig. 25). 

| These fruits, both in size and structure, bear a striking 
@ resemblance to the “stones” of the drupaceous fruits of 

Cy Curtisia faginea. This resemblance is supported by the 

identity of cell structure in the outer wall. Tentatively 

Fic. 25.— these bodies may beregarded as being the fruits of thattree. 
Sea ‘““Two-chambered Fruits.’’—These are the commonest 

of the fruit bodies and are locally abundant in the rock. 
They are smooth ovoid bodies 8-12 mm. long and 6-8 mm. across. 
Externally they are slightly pointed at one end and blunt at the 
other. In many there is a groove running round the fruit longi- 
tudinally which gives an appearance of division into two. Internally 
each is divided into two chambers by a partition which ran longi- 
tudinally down the middle. The groove is at the line of junction of 
the partition and the outside wall. 

The partition is complete, but is much thinner at the point and is 
often broken down there. 

The majority of these fruits appear quite empty or are filled with 
amorphous material. A few, however, have been found which have 
contents. In each chamber is a body flattened along the septum 
but convex on the outer side. Near the base of the flat surface is a 
rounded area which forms the point of attachment. Hach of these 
internal bodies has a rugose surface. 

In one piece of the hard rock there is a two-chambered fruit of 
a different kind. This is a rounded, rather flattened body about 
4 by 3 mm. and 3 mm. thick. Broken across it shows a thick wall 
composed of uniform parenchyma-like cells. There is a partition 
dividing the interior into two unequal portions. The interior is 
filled with chalcedony and does not show any certain structure. 


Fossil Plants from Fort Grey near East London. 89 


MiscELLANEOUS ANGIOSPERM STRUCTURES. 


“Smooth Fruits.’”’-—These occur in the same rock with the “two- 
chambered fruits” described above, but are much less abundant. 
They are nearly spherical and measure 9-12 by 8-9 mm. The ends 
are rounded, not at all pointed. They were quite smooth both 
externally and internally. On the inside there is a low ridge pro- 
jecting inwards. This ridge was not a complete partition, as there 
is no trace of it on the opposite side. The ridge runs longitudinally, 
but not quite straight. No contents have been found. 

“Flat Fruits.”—A few flattened structures, probably fruits, have 
been found. They are smooth and oval, 6-8 by 4—5 mm. in surface, 
and 2-3 mm. thick. When broken across they show a distinct thick 
outer wall enclosing a smooth body with a much thinner wall. No 
details of structure are available. 

“Smooth Seeds.’”,—From various parts of the rock and especially 
from the uppermost layers there have been obtained smooth ovoid 
bodies up to 2-5 mm. long and 2-3 mm. across. Some are nearly 
spherical, others are distinctly pointed at one or both ends. There 
is a considerable range in size and shape, and it is certain that more 
than one kind is represented. In some of the best preserved examples 
a low ridge runs obliquely up the surface on one side for about two- 
thirds the length. 

One specimen, perhaps, justifies further description. This is a 
smooth ovoid body, 4 by 2-5 mm. The specimen has been partly 
broken, and the break reveals a distinct double wall. The outer and 
inner walls are connected by numerous fine strands that run straight 
across between them. 

“‘ Ribbed Seeds.’’—These are ovoid bodies 3-5 by 2-3 mm., in which 
the surface is marked by longitudinal grooves or ridges. These occur 
as impressions. They were bluntly pointed at one end. On the 
surface the grooves are much wider than the intervening ridges. 

In some the grooves and ridges are simple, in others they are 


‘interrupted or complicated in various ways. 


“Prickly Seeds.’”’—Three examples have been found of ovoid or 
nearly spherical bodies about 2-3 mm. in diameter which have an 
unridged surface covered by sharp projecting points. In two of the 
three specimens these points are irregularly arranged—in the third 
they are in lines. 

Fungi.—Hyphae of a fungus have been found in more than one of 
the sections in the tissues of other plants. They have been noted in 


90 Annals of the South African Museum. 


the Gymnosperm wood and in the roots of ferns and in various 
unidentifiable fragments. No spores or spore-producing structures 
have been found. 

Structures of Uncertain Affinity.—In the rocks are numbers of plant 
fragments that cannot at present be assigned to any definite group. 
Some of these are very fragmentary, but some are quite definite and 
distinctive. Of these uncertain fossils by far the most abundant 
and important are leaves that for want of any means of identification 
are referred to as “Ribbed Leaves” on account of their general 
appearance. 

“Ribbed Leaves.”’—Locally in the uppermost bed there are large 
numbers of fragments of parallel-sided leaves which are marked by 
prominent longitudinal ribs. These occur in a poor state of pre- 
servation or as impressions. In one part of the rock they occur in 
very large numbers; masses of these fragments are found together 
in layers. The leaves themselves run in all directions. The rock 
here is hard and does not split along any definite planes, so that it 
has not been possible to obtain really satisfactory specimens. Not 
one example has been seen in which a complete leaf is present. The 
largest piece is 9cm.in length. In width the pieces are very uniform, 
about 2—2:5 cm. 

These leaves taper at both ends: at the base rather gradually to 
about 0:75 cm.; more abruptly at the tip, which is rounded but 
acute. 

One specimen has been found which shows what may be the 
attachment of these leaves to an axis. In this there is a much 
flattened axial structure, 13-5 cm. long, with the basal parts of 
leaves attached at either side. There are three leaves on one side 
and five on the other. The leaves are narrowed to their attachment, 
but show no sign of any sheath. The leaves are approximately 
opposite on the axis. The latter tapers: it is 2 cm. across at one 
end and only 1-25 at the other. 

In the leaves themselves the ribs are much more prominent on 
one side than the other: the ribbed side is presumably the lower. 

The flat surface is without stomata. It has an epidermis of 
elongated lozenge-shaped cells which appear uniform all over. The 
under surface has numerous stomata arranged in bands in the grooves: 
there are none on the ridges. The epidermis is made up of elongated 
lozenge-shaped cells; narrow on the ridges and broader where the 
stomata occur. The stomata are sunk somewhat below the surface. 
Each depression is surrounded by 4-6 cells: examples with four, five, 


dl 


Fossil Plants from Fort Grey near East London. 91 


and six cells have been found on one leaf. There are two cells much 
smaller than those of the general epidermis adjacent to the guard 
cells which are accessory cells (fig. 26). 

The stomata are in lines, but not regularly so. 

Sections show a rather thick leaf. The epidermal cells project 
somewhat. Subepidermal fibres occur along the flat upper surface 
and on the ridges on the lower surface. Of the interior about two- 
thirds is occupied by palisade cells. In none of the sections is any 
definite vascular tissue recognisable. Areas of structureless material 
represent the positions of veins. 

The axis which has probably leaves in attachment is very much 
compressed and has little definite 
structure preserved. It appears to be 
made up of strands running in a mat- 
~rix, but no certain cell outlines can be 
made out. 

With the material available it is not 
possible to assign these leaves to any 
group. The specimen referred to in 
which attachments occur may be a 


. ; Fic. 26.—Epidermis of ribbed 
portion of a stem bearing leaves, or ibe TL. 


a piece of a large pinnate leaf. A pro- 

longed search failed to give any further specimens that could afford 
evidence that might decide this point. One specimen with impressions 
only shows an axis with three leaf bases. There is no indication 
of any kind of bud on this: the base of the leaf passes into the 
axis gradually on the lower side; on the upper side the outline 
curves upwards just at the point of attachment, giving a rounded 
angle. While the specimen is incomplete and entirely without cell 
structure it seems to support the possibility of these leaves being 
pinnae of a compound leaf. 

On the other hand, the large numbers of leaf fragments and the 
great rarity of portions attached to any form of axis is not easily 
explained if this is the case. 

The lack of recognisable vascular structures renders any attempt 
at identification a matter of mere speculation. 

The general appearance and shape are suggestive of monocotyledons, 
but the absence of a sheath seems against associating them with that 
group. The epidermal structure seems to eliminate the possibility 
of a cycadean affinity. 

The large mass of these leaves was found in close proximity to a 


92 Annals of the South African Museum. 


stem which had wood of the type identified as Podocarpus, but there 
is nothing except their positions in the rock to suggest any sort of 
connection. 

Miscellaneous Structures.—(a) In two sections are fragments that 
represent portions of the wall of a rounded body such as a spore 
or pollen grain. These are thick bands with, on the outside, a series 
of sharp projecting points. They are made up of continuous material 
and not of separate cells (fig. 27). 

(6) In close proximity to one of the above is a small portion of 
tissue with cell structure. This consists of 2-3 layers of parenchyma 
cells connected to a layer of large columnar cells with walls very 

much thickened all round. The fragment is prob- 
ably a bit of a seed coat. 
(c) In one section there occurs an oval body about 
‘ \ 1:5 mm. along the longer axis. This has a somewhat 
irregular wall through which runs a band of thickened 
columnar cells. The interior is hollow. 


Fic. 27.—Spore- Various other scraps and fragments have been 
like structure. 


x 42. found which it is impossible to assign to any group 
of plants. Most of these are so uncertain in nature 
that no descriptions are needed. 

General Considerations.—The fossil plants described above without 
doubt represent the remains of a land flora. Not one of the structures 
is of a nature to suggest an aquatic or even a marsh environment. The 
roots with their wide large-celled cortex might possibly at first sight 
seem to be associated with moist conditions, but it has been shown 
that the cortex here is not in the original condition and that the 
existing plants that give the most probable comparison are certainly 
plants of dry land. 

The distribution of the roots in the rock would point to their 
preservation wm situ. This is supported also by the positions of a 
large number of the gymnosperm axes which has already been noted. 
In the quarry at the south extremity of the rocks gymnosperm axes 
pass down without break from the silicious rock into extremely 
weathered dolerite below. 

As regards the original conditions, this preservation im situ, and 
especially the penetration of the tree-roots into the dolerite, com- 
pletely rules out any possibility of transportation and redeposit. 
The origin that is most likely, and which explains most readily the 
nature and distribution of the fossils, is that the soil in which the 
plants were living has been impregnated with silica and so con- 


Fossil Plants from Fort Grey near East London. 93 


solidated. Preservation of the soil would give the explanation for 
the great abundance of roots and the scarcity and fragmentary 
condition of the leaves. 

The gymnosperm wood also seems to fit in with the soil nature of 
the rocks. The largest portions, which would be the aerial stems, 
are prostrate and often in short pieces, while those im situ, which are 
roots, are smaller. 

A soil would also explain the apparent complete absence of animal 
fossils: such animals as were living in the soil would leave no trace 
behind. 

The woody gymnosperms, and especially Podocarpus, are of quite 
general occurrence in the rocks. They occur in all the layers, and 
are not noticeably more abundant in one than another. The roots, 
too, are generally distributed through the middle and upper layers. 
In the basal beds the absence of preserved structure, except in wood, 
renders it uncertain whether the numerous casts and tracks which 
occur belong to these roots or to rootlets of gymnospermous trees. 
On the other hand, the other plants seem very much more local in 
their distribution. The cupressoid gymnosperm twigs, which are 
tentatively associated with the less abundant type of wood, are only 
found in portions of limited extent in the uppermost beds and in the 
upper part of the hard flinty rock. 

Local distribution is still more evident in the cases of the soleno- 
stelic ferns, the scapose monocotyledons, and the ribbed leaves. 
Hach of these is abundant where it is found, but the area occupied is 
very limited. The protostelic ferns, though never abundant, are more 
widely spread through the rocks, especially in the upper portions. 

Relations of Fossils.—In these rocks the gymnosperm wood, even in 
the most favourable cases, shows signs of alteration and of decay 
having taken place before silicification. Specimens have been ex- 
amined from all the beds of rock, and none has been found in which 
the softer portions, such as the bark, have been preserved. Bark is 
completely absent in all the larger pieces; and even in small ones, if 
present, is in a poor state of preservation. In many examples the 
wood itself shows signs of decay: the outer surface in some is eroded 
and irregular: radial cracks have been noted above. In one large 
piece the centre was not closely connected to the outer parts of the 
wood. Further, the generally found ghostlike nature of the pits 
on the cell walls and the very poor preservation of the parenchyma 
of the wood and medullary rays point to decay before impregnation 
and fossilisation. 

WiOl xoxox. PART di. 12 


94 Annals of the South African Museum. 


In contrast to this condition of the wood, the roots frequently show 
excellent preservation, not only of the harder parts but also of the 
much less resistant thin-walled cells of the cortex. Alteration has 
taken place, but it is of a different kind, occurring in preservation 
and not previous to it. 

It may be noted in this connection that gymnosperm roots of small 
size, which may occur in the same rock with these other roots, are 
not nearly so well preserved as the latter. 

Several examples have been found in which portions of wood have 
been penetrated by roots. These may make their way along cracks 
or even penetrate the wood itself. Sections of such root-penetrated 
wood show excellent preservation in the roots and a good deal of 
alteration in the wood. 

These various results strongly suggest that changes in the vege- 
tation were occurring previous to fossilisation. The wood decay 
indicates that the trees were already dead, while the state of the roots 
and their penetration into wood show a later period of activity. 
The cessation of activity of the trees previous to fossilisation is 
supported by the prostrate position and often short lengths of the 
stem portions and the frequent presence in the uppermost beds of 
small isolated pieces of woody stems. 

Taking all the available data into account, it seems probable that 
a gymnospermous forest, mainly composed of Podocarpus, existed 
at one time on this site, but was destroyed. On destruction the 
underground portions remained in position and were eventually pre- 
served before decay had proceeded too far. The upper parts fell 
and broke up to a large extent. On destruction of the forest, 
herbaceous plants became exceedingly abundant, as shown by the 
great numbers of their roots that have been preserved. 

The cause of the destruction of the forest is a matter of speculation. 
There is no evidence for subsidence and waterlogging of the soil 
which might be an explanation. Another factor that might have 
operated is fire. Trees killed by fire may remain in position for a 
long period, and direct regeneration may not occur. It is conceivable 
that the local abundance of ferns is an indicator of fire. At the present 
time Gleichenia polypodioides may become very abundant locally on 
burned areas of forest at Knysna (cf. Phillips, “Forest Succession and 
Ecology in the Knysna Region,” Bot. Survey S. Afr., xiv, p. 236, 
1931). ) 

Age of the Rocks.—The fossil plants that occur here, so far as they 
can be identified at all, are such as would be expected in a fairly 


Fossil Plants from Fort Grey near East London. 95 


recent rock. All are types that might be in existence at the present 
time. 

The fossil flora shows a striking difference to that existing in the 
locality at the present time in the dominance of gymnosperms and 
the relative paucity of angiosperms. The latter are both less in 
abundance and smaller in size in the fossil flora. 

Whether this difference represents a real difference in the flora as a 
whole that existed, or is the result of the preservation of a local 
community of plants, can only be settled by the study of contem- 
porary deposits over a wider area. 

At the present day Podocarpus occurs in the region, but at this 
low altitude only as isolated individuals in communities definitely 
dominated by angiosperms. Continuous forests of this tree, even 
of small extent, do not occur. 

The differences in the fossil flora as compared with the existing one 
at this altitude point to a not very recent date for the rocks. On the 
evidence available a Tertiary age is the most probable, though the 
position in that epoch must await the accumulation of further data. 


SUMMARY. 


1. Near Fort Grey is a local surface quartzite containing plant 
fossils. The plant remains are numerous, but fragmentary. 

2. The quartzites overlie dolerites. 

3. The most abundant fossils are woody stems and roots of a type 
that is identified with Podocarpus. Other gymnosperms are more 
like Widdringtomia. 

Leaves, twigs, and possible seeds can be associated also with these 
trees. 

4. Ferns occur rather locally. Some specimens are identified with 
Gleichenia. Other ferns are represented by solenostelic and proto- 
stelic stems, petioles, roots, and scraps of leaves and sporangia. 

5. Both monocotyledons and dicotyledons occur, the former as 
rhizomes, aerial stems, and roots; the latter as roots, small woody 
stems, scraps of leaves, and fruits and seeds. Curtisia has been 
identified by fruits. 

The roots are shown to have undergone shrinkage, and some experi- 
ments to demonstrate this are described. 

6. Some large ribbed leaves of uncertain affinity are described. 


96 Annals of the South African Museum. 


7. The general distribution of the plants in the rocks is discussed, 
and it is considered that the rock represents a solidified soil. 

8. The gymnosperm wood is in less good condition than the roots 
and represents an earlier phase of the vegetation. 

9. The fossils are part of a land flora. 

10. The rocks are of Tertiary age. 


( 97 ) 


3. On some Karroo Fishes from Central Africa—By 8. H. Hauveuton, 
B.A., D.Sc., Hon. Keeper of Palaeontological Collections. 


(With Plates XXIX-XXXI and 2 Text-figures.) 


Ischnolepis bancrofti gen. et sp. nov. 
(Pl. X XIX.) 


Co-types.—Two almost complete fish on slab No. 9338 in collection 
South African Museum; specimen No. 9339 in same collection. 

Locality—Lunsempfwa Valley, N. Rhodesia (lower portion); 
collected by Dr. Davis, and presented by Dr. J. A. Bancroft. Dis- 
covered in “a brown ferruginous shale with a maximum thickness 
so far found exposed of from 3 to 4 feet. Above this brown shale is 
a thin bed of a tough calcareous shale showing a peculiar cone-like 
structure” (Davis). Associated with fossil plants and Palaestheria. 

Horizon.—Probably Upper Beaufort or Lower Stormberg. 

Generic Characters.—Body fusiform. Mandibular suspensorium 
oblique, gape moderately wide. Dentition consisting of sharp, well- 
spaced, delicate, anteriorly directed teeth in front part of mandible 
and maxilla, and sharp, robust, more closely packed teeth in the 
hinder two-thirds. Opercular apparatus relatively small; inter- 
operculum well developed. One X-bone present in front of pre- 
operculum. Fins with fulcra, forked distally, articulated except in 
pectoral; pectoral fin relatively short-based; pelvic fin long-based; 
dorsal fin remote, opposed to anal, rather high in front, with a 
moderately elongated base-line; anal fin much extended, high and 
acuminate in front, its base reaching almost to the lower caudal 
fork; caudal fin almost equilobate, body lobe not extending to distal 
end, and fairly blunt-ended. Scales small, rhomboidal, arranged in 
numerous rows passing slightly sigmoidally backwards from dorsal 
to ventral margins of body. Lateral line prominent, surrounded by 
fine tubes. 

Specific Characters.—As for the genus. Fish small, up to 75 mm. 
in length. Head occupies about one-fifth of the entire length, greatest 
depth of body a little more than length of head, and in neighbourhood 
of pelvic fin. 


98 Annals of the South African Museum. 


Description.—The details of structure are obtained from the study 
of several specimens and a number of body and head fragments; no 
specimen is complete enough to show all the features. The arrange- 
ment of the bones of the head in the postorbital region is Palaeoniscid. 
A number of details can be obtained from a specimen that gives an 
inner view of the bones of this region. 

The maxilla has a low suborbital portion and a higher and long 
postorbital part; the latter has a regularly curved upper border, and 


/ co = ie WEAK 
= Ss wy SSS 


Text-Fic. 1.—Ischnolepis bancrofti, Htn. 
Restoration of body, drawn by Dr. K. H. Barnard. 


has no pronounced upper posterior angle such as is seen in Palaeonis- 
cus. The upper portion of the postorbital part of the bone is excavate 
on its inner surface, while the lower half is furnished with a series of 
small pits, which are elongated in an antero-posterior direction. 

The mandible is rather weak, with a pointed anterior end. Its 
inner surface is apparently ornamented with irregular longitudinal 
grooves and ridges. It is not possible to determine whether it and the 
maxilla carry one or two rows of teeth. In the hinder two-thirds of 
the dentigerous borders the sharp-pointed but robust mandibular 
and maxillary teeth interdigitate closely; the anterior sharp-pointed 
slender teeth are more widely spaced. 

The preoperculum is of the usual Palaeoniscid shape, with a narrow 
vertical posterior part forming the hinder boundary of the maxilla, 
and a deeper upper part lying above the maxilla. The front border 
of the preoperculum is excavate for the reception of a suborbital 
element. The long posterior margin meets the suboperculum, the 


— 


On some Karroo Fishes from Central Africa. 33) 


operculum and, superiorly, the supratemporal, and is convex. The bone 
is pierced, along its posterior border, by the preopercular sensory canal. 

The operculum is a relatively small, somewhat triangular, plate, 
with a rounded ventral apex and a dorsal width nearly equal to its 
height. The upper anterior corner is of uncertain shape, but it is 
apparently separated from the preopercular by a small triangular 
ventral prolongation of the supratemporal, which may be a separate 
element corresponding to Tra- 
quair’s bone X. 

The suboperculum is larger 
than the operculum, and is higher 
than wide. Its anterior border 


X Sup.Tem, 


is concave, its posterior and lower Y 

borders convex. The upper Y 

border is slightly excavate, so eS Uf 

that there is a loose junction P 

between the bone and the oper- GEN 

culum. The upper anterior corner Ses 

is acute-angled. TExt-FiG. 2.—Ischnolepis bancrofti, Htn. 
Ventrally to the suboperculum Restoration of posterior part of 


: 3 ; skull, lateral view. 
is a smaller reniform bone which 


probably belongs to the opercular series, and is here interpreted as an 
interoperculum. It lies behind the hinder end of the mandible, and 
has the same internal surface ornamentation of small circular pits 
as the suboperculum and operculum. The branchiostegal rays 
undoubtedly passed forward from the lower border of this bone; but 
no specimen shows them in contact with the bone, although their 
presence is confirmed by one fragmentary example which is seen in 
ventral view and which presents, on either side of a median triangular 
bony mass, a series of four or five overlapping branchiostegal plates 
lying mesial to the mandible. 

In his recent paper on “Fossil Fishes from the Karroo System,”’ 
Brough (P.Z.8., 1931, p. 234) figures the bone which is here described 
as an interoperculum, but describes it as probably the modified first 
branchiostegal ray, stating that it “does not correspond to the inter- 
operculum.” He also asserts the absence of branchiostegal rays in 
some Catopterid genera, and thinks that they may be absent in all. 
The specimens from Northern Rhodesia described here as one species 
usually show no trace of such rays; and their presence is only con- 
firmed by one fragmentary example which is, fortunately, seen in 
ventral view. 


100 Annals of the South African Museum. 


The ornamentation of the opercular bones on the external surface 
presumably consisted of small, rather scattered, low tubercles. 

The limits of the membrane bones of the cranial roof are difficult 
to decipher exactly, but they can be drawn approximately from the 
positions of the sensory canals that penetrate them, which are pre- 
served as raised tubes on the inner face of the skull. 

Above the preopercular the lateral canal runs longitudinally through 
the supratemporal, which has apparently a ventral triangular portion 
wedged in between the opercular and preopercular. The supra- 
temporal (““supratemporo-intertemporal”’ of Stensi6) has a somewhat 
digitated suture with the parietal and a straight suture with the 
tabular (“extrascapular” of Stensid), which lies behind it. This 
latter carries the transverse tabular canal. The lateral canal cannot 
be traced in the tabular, which is a short broad bone; but its curved 
course through the presumed post-temporal (“suprascapular” of 
Stensi6) can be followed, and its continuation through the upper 
posterior corner of the supracleithrum to the lateral line of the 
body. 

The shoulder-girdle is not fully displayed in any one specimen, but 
the cleithral and supracleithral are typically Palaeoniscid. The 
former is a bent pillar-like bone ornamented with longitudinal ridges; 
the supracleithral is an expanded plate with a strengthened anterior 
border. In front of the ventral prolongation of the cleithral, and 
partly enwrapping it, is a long, pointed triangular clavicle which, in 
one specimen, seems to be fused with its neighbour in the middle 
line. 

The pectoral fin is small and low down on the body. The fin-rays, 
which are possibly about 15 in number, are forked distally, and seem 
to be unjointed throughout their length. There are well-developed 
small fulera. 

The pelvic fin is considerably larger than the pectoral, with about 
20 rays, and is in front of the middle of the body. The rays are 
articulated near their bases, but apparently not distally. The rays 
are forked distally, and fulcra are present. 

The anal fin has a long base, and in the anterior part is relatively 
high. The rays number about 50, but the baseosts are less than half 
this in number. The rays are forked and, at least in the anterior 
half of the fin, articulated. 

The base of the dorsal fin is about half the length of that of the anal, 
and the number of fin-rays is about 20. In front of the fin there are 
a few enlarged pointed dorsal ridge-scales, and prominent fulcra are 


see 


On some Karroo Fishes from Central Africa. 101 


developed. The front of the fin is opposite the front of the anal. 
The fin-rays are forked distally, and articulated throughout their 
length. The baseosts number only about half the fin-rays. 

The caudal fin is deep and fairly deeply cleft, with the lobes about 
equal to one another and not excessively elongated. The body lobe 
is fairly long and somewhat bluntly pointed. All the rays of the fin 
are articulated and branched. At the base of the upper lobe are a 
number of elongated dorsal ridge-scales, and the anterior border of 
each lobe is furnished with fulcra. 

A typical fish has an estimated length of 75 mm., a greatest depth of 
body (at pelvic fin) of 16-5 mm., and the length of base of anal fin 
17mm. The body has about 50 transverse rows of scales, which are 
small and very thin, and are apparently ornamented with two or three 
irregular longitudinal ridges. The lateral line of the body is enclosed 
in small tubular bones lying in a continuous longitudinal row, and 
continues backwards to the base of the caudal fin opposite to the point 
of bifurcation. 

This fish, while bearing some resemblance to the Upper Beaufort 
genus Helichthys, does not seem to fall within the limits of any of the 
hitherto described Catopterid genera. The combination of characters 
presented by the head, body, fins, and scales—the small size, the 
smallness of the scales, the relative positions of the dorsal and long- 
based anal fins, and the probable presence of a well-defined inter- 
operculum—give it a very distinctive appearance; one or more of 
these characters can be paralleled in one or another of the genera 
which have an oblique mandibular suspensorium, but they are not 
all found in combination in any of the known forms. The description 
of Urolepis given by de Alessandri almost fits the specimens under 
discussion; but his photographs of typical examples of that genus 
display distinct points of difference. It thus becomes necessary to 
found a new genus for these fishes, which can be known as Ischno- 
lepis bancrofti gen. et sp. nov., the tribal name being given in honour 
of Dr. J. Bancroft, to whom the opportunity of studying these speci- 
mens is due. For the accompanying drawing of one of the co-types 
I am indebted to Dr. K. H. Barnard. 


Pygopterus (*%) sp. 


A broken slab and partial counterslab display part of the body of a 
small-scaled fish in which the axial skeleton is well seen. Although 
the anal fin is not preserved and only a portion of the dorsal fin 


102 Annals of the South African Museum. 


remains, the latter was certainly small compared with the anal and 
did not arise in front of it. 

Above the notochord there is a series of neural arches which 
diminish in height posteriorly, and whose height in the middle of the 
specimen is slightly less than the diameter of the notochord. The 
head of each arch is slightly dilated, and the arches are inclined back- 
wards at about 45° to the axis of the notochord. Above each arch is 
an elongate slender neural spine whose proximal end only is dilated. 
The dorsal fin is supported by two rows of interspinous bodies, 12 or 
13 in each row. Of these the axonosts are longer than the baseosts; 
both sets are more elongate than in “Oxygnathus” brown. Both 
sets are slender, hour-glass shaped bones, and the length diminishes 
rapidly in the posterior half. The lepidotrichia of the dorsal fin are 
far more numerous than these endoskeletal radials. 

Haemal arches occur from front to back of the specimen. They 
are longer than the neural arches, considerably swollen proximally, 
and only very slightly expanded distally. Hach articulates with a 
long slender haemal spine. The anal fin is supported by a number of 
elongate endoskeletal radials arranged in a single row, each of which 
is dilated distally. 

The relative sizes of the endoskeletal elements in this fragment are 
sufficient to differentiate the form from Oxygnathus brownt, to which it 
approximates in size. The body scales were obviously small and 
thin; and the position of the dorsal fin with respect to the anal renders 
it possible that the fragment falls within the limits of Pygopterus. 

This description is based on specimen No. 9340 in collection South 
African Museum, from the lower part of the Lunsempfwa Valley, 
N. Rhodesia. The slab and counter-slab were collected by Dr. Davis 
from the beds that yielded Ischnolepis bancroftt, and were presented 
by Dr. J. A. Bancroft. 


Cf. Atherstonia sp. 
(Pls. ROXUX. > XO IE) 


Three specimens, obtained by Dr. Davis from the beds which 
yielded Ischnolepis bancroftt in the lower part of the Lunsempfwa 
Valley, North Rhodesia, give evidence of the presence of a large 
Palaeoniscid fish, although they are not complete enough to be 
generically identifiable. Each will be briefly described. 

Specimen 9351 (in coll. S. Afr. Mus.). 

Contains part of the head and pectoral fin. Operculum high and 
narrow, ornamented with irregular, closely set ridges and elongate 


On some Karroo Fishes from Central Africa. 103 


tubercles. Suboperculum very much smaller, subpentagonal in shape, 
as wide as high, ornamented with a central mass of rounded tubercles 
surrounded by irregularly curving ridges and elongate tubercles. 

Branchiostegal rays numerous—23 being preserved ; rays much 
wider than high. 

Pre-operculum long and well developed, situated very obliquely. 
Bone slightly bent, rodlike posteriorly and expanded anteriorly. 

Maxilla large, presumably Palaeoniscid in form. 

Dentition (seen on mandible only) consists of stout conical teeth of 
two sizes, larger and smaller, rather widely spaced. 

Pectoral fin incomplete; large, consisting of about 21 rays, of 
which most show dichotomous forking; fulcra numerous, but small; 
articulation doubtfully present in anterior rays. 

Specimen 9350 (in coll. S. Afr. Mus.). 

Slab and partial counter-slab, showing most of the body, lacking 
the tail and dorsal fin, together with the badly preserved posterior 
part of the head, and the pelvic and anal fins. Tentatively, it is 
considered to belong to the same species as the foregoing, although 
it is rather smaller. Body fusiform. Depth of body about half the 
length from posterior edge of clavicle to root of tail. Scales rather 
small with non-denticulated posterior borders, and ornamented with 
irregular, rather weak, forked, and anastomosing ridges. Continuous 
series of dorsal ridge-scales. 

Pelvic fin rather closer to anal than to pectoral; rays nearly 30 in 
number, articulated, distally bifurcated; anterior rays longer than 
fin-base. 

Anal fin long-based; rays numerous, articulated. 

Position of dorsal fin not exactly determinable, but probably 
opposite to, or slightly in advance of, anal. It is certainly not behind 
the anal. 

Specimen 9353 (in coll. S. Afr. Mus.). 

The greater part of a tail, which probably belongs to this species. 
Tail strong and forked, upper and lower lobes about equal. Rays 
strong, jointed throughout their length, dichotomously forked. 
Dorsal lobe has ridge-scales of medium strength; ventral lobe has 
rather small fulcra. There are shown five strong haemal spines 
forming a support for the anterior part of the ventral lobe. The body 
squamation extends for a considerable distance up the dorsal lobe, 
and ends in a sharp point. 

On account of the uncertainty concerning the position of the 
dorsal fin, it is not clear whether these specimens belong to that 


104 Annals of the South African Museum. 


group of the Palaeoniscidae that contains the genera Acrolepis, 
Gyrolepis, Atherstona, Myriolepis, and Oxygnathus, or to that con- 
taining Pygopterus and Urolepis. It differs from Acrolepis in the 
length of base of the anal fin and in the size of the scales. It agrees 
with Gyrolepis, particularly in the relative narrowness and depth 
of the operculum and in the nature of the fins; it is larger than most 
of the described species of that genus, and its scales are smaller and 
not so deeply overlapping. The dorsal ridge-scales are not so pro- 
nounced as in Atherstonia, and the body-scales are smaller; the 
ornamentation on the scales is somewhat similar. Myriolepis differs 
in the possession of smaller scales and in having a short-based 
anal fin. In Oxygnathus, too, the scales are smaller and the body 
is more elongate. 

The generic position of this form is thus rather doubtful. Of 
those cited, Acrolepis, Atherstonia, and “‘Oxygnathus”’ (Broom) are 
known from deposits in South Africa and Madagascar; and it would 
seem best temporarily to designate the forms under consideration 
simply as Cf. Atherstonia sp., recognising that there are certain points 
of difference that mark them off specifically at least from other 
members of the genus. 


Ann. S. Afr. Mus., Vol. XX XI. Plate X XIX. 


ISCHNOLEPIS BANCROFTI Gen. et Sp. N. 
Co-types: Two fish on Slab 9338. 


Neill & Co., Ltd. 


ee ee cm a Re RR oe FE a LN Ae NTN AD AEN GI IT LT LT OLLI LEELA OLD: TLL: Si EO ALLL ALG AL LOL ee ee ed 


maf Dade 
Se 


a a ai wee 
f y - ae = 


Plate XXX. 


Ann. S. Afr. Mus., Vol. XX XI. 


‘0SE6 GRIS “USY Teed 
dg VINOLSYUAHLV 


fO 


Neill & Co., Ltd. 


| 


Ann. S. Afr. Mus., Vol. XX XI. ¢ Plate XX XI. 


eres Ojo: 


Max. 


Mand. 


Branchial 
Rays. 


Cf. ATHERSTONIA 


a. Slab 9351. b. Slab 9353. 
Neill & Co., Ltd. 


( 105 ) 


4. The Phosphatic Nodules of the Agulhas Bank. (A Study of Sub- 
marine Geology.)—By L. Cayrux, Membre de l'Institut, 
Professor at the Collége de France, Paris. 


(Translated into English by Dr. 8. H. Haveuron.) 
(With Plates XX XII-XXXV.) 


THE cruise of the Challenger was responsible for the discovery of 
phosphate of lime among the marine sediments which are accumu- 
lating in our day. According to J. Murray, A. F. Renard,* and 
L. W. Collet, ¢ the Challenger dredged nodules of phosphate of lime 
on the Agulhas Bank, south of the Cape of Good Hope, on the eastern 
sides of Japan and of Australia, on the coasts of the Argentine 
Republic, between the Falkland Isles and the estuary of the Plata, 
to the west of Chile, and on the coast of Spain. Further, phosphatic 
nodules have been recovered by the Blake { at numerous points 
on the coast of the United States between Florida Strait and Cape 
Hatteras, and in the North Pacific. Finally, the German cruises of 
the Gazelle and of the Valdiia,§ not to mention the important 
researches of the Department of Agriculture of the Cape, have 
dredged much phosphatic material on the Agulhas Bank, previously 
explored by the Challenger. In our present state of knowledge it 
can be said that the three great meridional oceans have each yielded 
phosphatic concretions. 

J. Murray and A. F. Renard (1891) and, above all, L. W. Collet 
(1905 and 1908) have given detailed descriptions of the nodules 


* J. Murray and A. F. Renard, “ Deep-Sea Deposits” (Report of the Scientific 
Results of the exploring Voyage of H.M.S. Challenger), 1891, Phosphatic Con- 
cretions, pp. 391-400, pl. xx. 

+ L. W. Collet, “Les concrétions phosphatées de l Agulhas-Bank,” Proc. Roy. 
Soc. Edin., vol. xxv, 1905, pp. 862-893; “‘Les dépots marins, 1908,’ Les con- 
crétions phosphatées, pp. 194-213. 

{ J. Murray, “Report on the Specimens of Bottom Deposits (Cruise of the 
Blake in the Atlantic, 1877-80), Bull. Mus. Comp. Zoology, vol. xii, 1885-86, 
pp. 37-61; A. Agassiz, Three Cruises of the Blake, vol. i, 1888, pp. 275-276, and 
vol. ii, 1888, pp. 281-282. 

§ Sir John Murray and Professor E. Philippi, Die Grundproben der Deutschen 
Tiefsee-Expedition, 1898-99, Bd. xx, 1908, pp. 181-187, pl. xxii. 


106 Annals of the South African Museum. 


recovered from the Agulhas Bank. Then Sir John Murray and 
Professor E. Philippi have devoted several pages to the concretions 
obtained by the Valdiia (1908). J. Murray had previously initiated 
these studies by a brief description (1885-86) of the phosphatic con- 
cretions dredged by the Blake on the coasts of South America. 

Although, however, phosphatic concretions have been dredged at 
a number of places, it would seem that, up to the present, but one 
true deposit has been discovered—that of the Agulhas Bank. To this 
reason for giving to the subject particular attention I add another 
—the very great interest which it presents from the theoretical 
standpoint. 

Thanks to the happy intervention of Dr. A. W. Rogers, formerly 
Director of the Geological Survey of the Union of South Africa, and 
to the kindness of Dr. EK. L. Gill, Director of the South African 
Museum, to whom I express my acknowledgments, I have had the 
good fortune to study some specimens of phosphatic nodules dredged, 
many years ago, by the Cape Government, under the direction of the 
late Dr. Gilchrist. The material examined came from nine different 
stations, ranging from 48 fathoms (88 m.) to 56-0 fathoms (1024 m.). 
A table of stations is given here.* 


caaaie Bearings of Station. ie ae Depth. 

i Lion’s Head, S. 82° E. 27 miles= 43-2Km.| 125 fathoms; 229 m. 
2 ve INSGa eee 345) sues) Odea: 154 a DBO nn 
3 Vasco da Gama Peak. New] hes.) =) s2e8-oiee 230 - aes 
4 Hang Berg, N. 2? E. 29) et g0 om ck 48 ae Shee 
5 Cape St. Blaize, N. x EH. 3? E. Yo ss) Gas 105 i 192i 
6 Cape Point, N. 44° E. 38 «yy = 60-89) 1315-100 576-731 m. 
7 ie N.E. 3 N. BO 5s Oa ee 560 ce 1024 
8 a N.E. x E. 3 EK. 2S eee 300 ao 549 ,, 
9 Lat. 36° 34’ S., long. 21° 32’ E. Me 240 af 439 ,, 
10 Off Vasco da Gama (specimen DOM not 166 . 304 ,, 


from L. W. Collet, P.F. Stat. 
XI). 


Note.—Lion’s Head is a part of Table Mountain, Cape. Vasco da Gama is a headland near 
Cape Point. Hang Berg is found at Cape Hangklip, on the E. coast of False Bay. 


To this material is added a nodule of great interest, dredged from a 
depth of 166 fathoms (304 metres) at station 10 (Vasco da Gama), 
which has been given me by Mr. L. W. Collet (see Map). 

* The map (Pl. XX XII) indicates the points of origin of my samples, numbered 


1 to 9. From my examination, I conclude that all the stations, except 4 and 5, 
are situated on the slopes S.W. of the Agulhas Bank. 


The Phosphatic Nodules of the Agulhas Bank. 107 


The nodules recovered by the Challenger came from two very dis- 
tinct regions. One group was obtained at 98 fathoms (179 m.) and at 
150 fathoms (274 m.) on the edge of the Agulhas Bank; the other at 
1900 fathoms (3475 m.), and thus in very deep water, nearly 100 miles 
S.E. of that Bank. 

Those which were sent to the Challenger office by Dr. J. D. F. 
Gilchrist, and studied by L. W. Collet, were taken from depths varying 
from 80 fathoms (146 m.) to 800 fathoms (1460 m.). 


External Characters of Nodules.—The specimens furnished by the 
Aculhas Bank and its neighbourhood have a striking morphology, 
seldom comparable with that of the pebbles of our Albian green- 
sands. Of a form extremely variable in detail, generally rounded 
but occasionally very angular (Pl. XX XIII, figs. 1-3), the nodules are 
irregular, mammiliform, ornamented with protuberances, even deeply 
indented, and—according to L. W. Collet—perforated by numerous 
holes. Certain of my specimens show a tubercular and scoriaceous 
appearance. One of them, cut through the middle, shows irregular 
funnel-shaped cavities reaching to the centre, and representing 
from a quarter to a third of the total volume. 

Representative in an unknown measure of the deposit, these nodules 
are divisible into two categories. The majority are characterised by 
an irregular form, and have a colour varying from a blackish-grey 
to pure black: the others are angular and brownish-yellow in tint. 
The former are dull, and the latter have a polished and varnished 
appearance, remarkably pronounced. In other words, the one group 
resembles concretions, while the other gives the idea of remanié rock 
fragments. These latter, which have on a single face cavities measur- 
ing 2 cm. in diameter and 1 cm. in depth, show further, on the same 
side, tiny perforations resembling cupules, 2 mm. broad, filled with 
glauconite. These differences of facies do not appear to me at all a 
function of the bathymetric conditions of the surroundings whence 
the nodules have been dredged. In this regard, it should be noted 
that the lightest colours (brownish-yellow) are found at depths of 
105 fathoms (192 m.) and of 230 fathoms (421 m.), while the blackish 
nodules have been dredged from 154 fathoms (281 m.) to 400 fathoms 
(731 m.) As to the scoriaceous pebbles, I can only say that the most 
typical of my specimens from this point of view came from the 
greatest depth, 560 fathoms (1024 m.). 

Many nodules carry incrustations of an organic nature, already 
recorded by L. W. Collet, who has identified corals, bryozoans, worms, 


108 Annals of the South African Museum. 


alcyonaria, sponges, and foraminifera with this interesting fact—which 
I have not been able to observe—that the incrustations can determine 
the positions of the concretions on the sea-floor and their degree of 
burial. One part, black and shiny, corresponds to the portion em- 
bedded in the sediment, and another—grey—to the incrustations 
defining the upper zone protruding from the mud. The incrustations 
visible on my samples, mostly due to bryozoa, are seen sometimes over 
the whole surface—proof that the nodules have been rolled in every 
direction—and sometimes over a part only. Several lack incrustations. 
According to J. Murray and A. F. Renard, the dimensions of the 
concretions vary mostly from 1 to 3 cm., and exceptionally reach 
4-6 cm. L. W. Collet states that they generally lie between 5 and 
10 cm. The largest, dredged from 111 fathoms (304 m.) at 25 km. 
from the coast (St. 10, Vasco da Gama), measures 23 x 16x12cm. The 
largest of my specimens does not exceed 7:5 cm. in greatest diameter. 
From the descriptions and figures given by L. W. Collet, Sir 
John Murray and Prof. E. Philippi, it is concluded that the nodules 
are either isolated or cemented to form a true conglomerate. As 
far as I can judge from the few indications furnished by the gangue of 
the nodules, it is formed of greensand more or less consolidated. The 
latter authors have figured a curious grouping of perforated nodules.* 
It must be noted here that, whatever their shape. the nodules have 
acquired the consistency shown by the hardest pebble of the Albian 
of the Paris Basin. 
Chemical Composition of Nodules—J. Murray and A. F. Renard 
have published the following analysis of concretions dredged at Station 
142, at 150 fathoms (274 m.), on the edge of the Agulhas Bank :— f 


PSOE : . 19-96 per cent. 
CO, Skt. SS 2 Eee 
SO, ye lesle Jducon’ leeagtiedae ere 
Si0, ; ; s6n sae 
Ghee ae ee) bo oe ee Oe 
MgO : : -,_ 0°6t Fe 
Fe,0, ’ ta eee 
ALOLG st Ae ure eae gaara 
Loss f : : ; ar 
Insoluble residue SVs: r 
95-89 ny 
Phosphate of lime . 2 4 S-Di 


* Sir John Murray and Professor E. Philippi, op. cit., pl. xxii, fig. 1. 
+ J. Murray and A. F. Renard, op. cit., pp. 392-393. 
{t Not estimated owing to an accident. 


The Phosphatic Nodules of the Agulhas Bank. 


An analysis of a nodule dredged by the Valdivia * gave: 


S10, . 26-70 per cent. 
Ca3(PO,). : ES Sc eee 
CacOs 7\- ST AO FOS y's: 
CasOn vr”. : : a WIEZ 6 anit, 
MeC@Osnerk « f : 4-67 i 
Fe,O, ' 8 es a 
Al,O3 ; : RRS ae 
102-38 


109 


Two analyses of yellow nodules, made by L. W. Collet, showed 
respectively 37-79 and 46-69 per cent. of phosphate of lime, and 
21-09 and 23-70 per cent. of Fe,O3;. Two other specimens studied 
by the same author, taken from one and the same station, yielded 


48-70 and 52-05 per cent. phosphate of lime. 


Three specimens which I have analysed, without estimating all the 
elements present,{ are characterised by the following compositions :— 


(10). 


(7). (8). 

SiO, LS SR ee 21-90 
LLOo a 3-00 3-00 
Hee eee, 6-25 6-25 
BeOmea ce 2 (Te lB 7-80 31-70 
MgO Radtaee Ayia ety cs yt 1-16 1-80 
POs Ree ils if nd 1125/1 dyallOo86 18-47 
a Yor 1-19 0:97 
ol hia 0:25 0-24 
Sar re POR a 8-14 4-20 
EO 2; yoy ee ae 5-75 7-20 

99-50 95-73 


13-60 
2-10 
6-30 

38-40 
1-44 

18-60 
1-35 
0-32 
7:93 
6-50 


96-54 


(7) Dredged at 560 fathoms (1024 m.), Cape Point. 
(8) Dredged at 300 fathoms (549 m.), Cape Point. 


(10) Dredged at 166 fathoms (304 m.), Vasco da Gama. 


* Sir J. Murray and Professor Philippi, op. cit., p. 185. 


t+ L. W. Collet, “Les concrétions phosphatées de  Agulhas Bank,”’ Proc. Roy. 


Soc. Edin., vol. xxv, 1905, pp. 868 and 870. 


{ In particular, the alkalis ought to figure in a higher proportion in the complete 
analyses of the nodules from Stations 8 and 10, on account of the frequency of 


occurrence of glauconite. 
VO. mex, PART I. 


13 


110 Annals of the South African Museum. 


The composition of these three nodules leads to the conclusion 
that the phosphate of lime is a fluocolophanite, a mineral to which 
A. Lacroix attributes a formula susceptible of variation : * 


z[(CaFl),Ca,(PO,),_] + yCaCO, + zH,0. 


From this composition it can be inferred that, from the chemical 
point of view, the nodules of the Agulhas Bank are allied to the 
group of phosphates of the ancient sedimentary series. 

These analyses show a remarkable similarity of content of phosphate 
of lime to those of J. Murray and A. F. Renard, and of Sir John 
Murray and EK. Philippi. The figures of L. W. Collet (values of 
46-69, 48-70, and 52-05 per cent. of phosphate of lime), whilst sensibly 
higher, do not invalidate the conclusion to be drawn from the other 
analyses, that the nodules of the Agulhas Bank fall within the category 
of poor phosphates. 

I would note, without stressing the point, that the figures of phos- 
phoric acid content are of the same order as those of the Albian 
greensands of the Paris Basin. 


MICROGRAPHIC STUDY. 


J. Murray and A. F. Renard have described and figured two types 
of concretions: (1) those found in the greensands of Station 142 of 
the Challenger, dredged at 150 fathoms (274 m.), compounded into 
aggregates, formed of minerals in the maximum proportion of 2/3, 
cemented by phos- 
phate of lime, with organisms absent; + (2) those found in the Globi- 
gerina ooze, and found at a depth of 1900 fathoms (3475 m.) (Station 
143), containing a preponderance of Foraminifera.{ The occurrence 
of the latter being independent of the Agulhas Bank, I leave them 
provisionally aside, but not without remarking on the extraordinary 
bathymetric dispersion of the nodules in question. 

I define here briefly six varieties of nodules, and afterwards shall 
consider them together. 

(1) Calcareo-phosphatic nodules, very quartzitic. [No. 1, 125 fathoms 
(229 m.), Lion’s Head, and No. 8, 300 fathoms (549 m.), Cape Point 
(Pl. XXXIV, fig. 1).]|—Blackish nodules, the richest in detrital matter 


of the whole series. Very uniform in size and scarcely ever exceeding 


9? 


comprising glauconite, quartz, and “silicates, 


* A. Lacroix, Minéralogie de la France, t. iv, 1890, p. 561. 
+ J. Murray and A. F. Renard, op. cit., pl. xx, fig. 1. 
{ J. Murray and A. F. Renard, op. cit., pl. xx, figs. 2-4. 


The Phosphatic Nodules of the Agulhas Bank. LTTE 


0-05 mm. in diameter (No. 8); the quartz grains (q) are unequally 
scattered in the restricted space of a section, of which they form a 
third, a half, or even more. On the average, they represent less than 
50 per cent. of the deposit. Contrary to the state of things in the 
nodules of the greensand, described by J. Murray and A. F. Renard, 
glauconite (g) may be rare, although one may count several dozen 
small grains in a preparation (No. 8). In the other nodules (No. 1), 
the mineral is thinly scattered. The fauna is characterised by the 
presence of some Globigerinae. The gangue is sparingly developed, 
and is calcareo-phosphatie. 

As can be seen, the rock is far from being a typical greensand. 

(2) Phosphatic or calcareo-phosphatic nodules, very glauconitic, with 
Globigerinae. [No. 4, 48 fathoms (88 m.), Hang Berg; No. 3, 230 
fathoms (421 m.); and No. 7, 560 fathoms (1024 m.), Cape Point. ]— 
The nodules from Stations 4 and 7, blackish and encrusted with 
glauconite, show indentations filled with this mineral. 

The grains of quartz and of glauconite alternate, but the glauconite 
has the greater relative volume. In the specimens examined, the 
quartz forms grains of variable diameter, seldom greater than 0-13 
mm., or elements whose diameter is always less than 0-10 mm. The 
glauconite is distributed in a most irregular manner, being totally 
absent sometimes, and at others forming by itself the greater part of 
the pebble. The Globigerinae are widely distributed, so much so 
that there is very little cement. 

Between this type of nodule and the preceding the differences are 
purely quantitative, and they are linked by transitional varieties. 

(3) Calcareo-phosphatic nodules with Bryozoa and Pulvinulinae. 
[No. 9, 240 fathoms (439 m.), 8. of Agulhas Bank.]—Nodules of very 
deep colour, in which the quartz, represented by grains measuring 
at the most 0:06 mm.—0-07 mm., is noticeably less. Glauconite is, 
on the average, less widely distributed or rare. Numerous very 
fragmentary remains of Bryozoa appear, of which at least a dozen 
can be counted in a section. The Globigerinae are considerably 
reduced in number, a reduction which is only partly compensated 
by a very marked increase in Pulvinulinae, which are generally char- 
acterised by a very thick test. The amount of gangue material is 
noticeably increased. The appearance is thus very distinct from that 
of the preceding types. 

(4) Calcareo-phosphatic nodules, quartzitic, and with abundant Glo- 
bigerinae. [No. 3, 230 fathoms (421 m.), Vasco da Gama (Pl. XXXIV, 
fig. 2).]—These nodules differ greatly from the preceding in their 


112 Annals of the South African Museum. 


buff-yellow colour. The quartz-grains (q) are more numerous, but do 
not show the same degree of frequency as in the nodules of the first 
two categories. Fairly variable, the maximum diameter reaches 0-13 
mm., while the minimum may be less than 0:05 mm. Glauconite 
is only an accessory mineral. The principal characteristic of this 
type is the abundance of Globigerinae and the presence of spicules 
of calcareous Sponges. Gangue material is very unimportant. 

The matrix of the nodules formed in this way is a passage-deposit 
to the true Globigerina mud. 

(5) Phosphatic and ferruginous nodules with Globigerinae. [No.5, 105 
fathoms (192 m.), Cape St. Blaize (Pl. XX XIV, fig. 3).]|—These nodules 
have an angular shape and brown colour, are almost free of quartz 
(0-05 mm.—0-10 mm.), very poor in glauconite, and particularly rich 
in Globigerinae. Their cement, which is both phosphatic and 
ferruginous, has preserved the organisms in an exceptionally beauti- 
ful manner. Numbers of these are riddled with perforations filled 
with oxide of iron. This type of nodule offers most affinity, of all 
the series examined, with the Globigerina ooze. 

(6) Phosphatic and ferruginous nodules with large benthic Fora- 
minifera and Bryozoa. [No. 6, 105 fathoms (192 m.), Cape St. Blaize 
(Pl. XXXIV, fig. 4).]—Although coming from the same station as the 
foregoing, these nodules have almost nothing in common with them 
(compare figs. 8 and 4). The organic remains, often so abundant 
that they touch one another, give to the thin section a very pro- 
nouncedly coarse appearance. They consist principally of the 
remains of benthic Foraminifera with extremely thickened tests, 
accompanied by accessory pelagic forms, especially Globigerinae, in a 
very small proportion. There are also fragments of bryozoan colonies 
and of parts of Echinoderms (plates and spines), both fairly frequent, 
and some fragments of the shells of Mollusca and of Brachiopoda, 
to the absolute exclusion of detrital minerals and of glauconite. In 
this chaotic complex, the pelagic Foraminifera may be arranged in 
rows. Most of the organisms are riddled with perforations which 
are filled with phosphate of lime and oxide of iron. Many are in- 
complete and penetrated by the matrix. Amorphous phosphate of 
_ lime and iron peroxide play a very important part both in the 
fossilisation of the test and in the infilling of the cavities. 

To these six categories I add another, very different from the 
preceding, and constituted of nodules of phosphatic greensand, whose 
characters will be discussed later (Pl. XX XV, fig. 3). 

From the examples thus passed in review, which possibly give an 


The Phosphatic Nodules of the Agulhas Bank. 113 


incomplete idea of the mode of occurrence, it is possible to judge 
of the variety of phosphate rocks represented in the deposit. These 
rocks include types obviously terrigenous, such as the calcareo-phos- 
phatic, very quartzitic, nodules and also the phosphatic and ferruginous 
nodules with Globigerinae, which resembles pelagic formations. Be- 
tween these two extremes is ranged a series of intermediate types 
whose terrigenous characters gradually disappear, step by step. 
The variety which is characterised by Bryozoa and numerous benthic 
Foraminifera appears to stand at the border of the complex. I see 
in it a representative of the numerous bottom deposits of organisms, 
formed in a terrigenous environment, which never figure in the 
classification of true marine sediments. 

In order to state fully the problem for solution, it is necessary to 
add that these phosphatic rocks have been dredged from a bottom 
of greensand, and that the specimens which most closely approach 
a Globigerina ooze—although deviating from the normal type by a 
very small proportion of quartz granules—have been recovered at a 
depth of only 105 fathoms (192 m.), 7.e. well outside the actual domain 
of the Globigerina oozes. 

As far as I know, no deposit of phosphate of lime in the older 
sedimentary series contains such a variety of types nor raises a 
problem of such importance. 


ANALYSIS OF THE CONSTITUENTS OF THE NODULES. 


The little that we know, up to now, of the constitution of the 
nodules scarcely allows us to suspect the great imterest which is 
attached to the elucidation of the general history of phosphatic 
concretions of all ages. Therefore I describe successively all the 
elements which go to form the nodules, and then discuss the structural 
details which give an unforeseen picture of the mode of origin. 

1. Minerals.—From one end to the other of the series analysed, 
quartz shows a great variation in frequency, since the grains may be 
predominant in the first variety and completely absent in the last. 
These variations entail only slight changes in the quantity of the 
elements, without showing a modification in shape. All are angular, 
with a mean diameter which is always less than 0-10 mm.; hence 
nearly all the grains would float in water that was only feebly agitated. 
The angular form of the grains, already noted by J. Murray and A. F. 
Renard, is a consequence of this. From our point of view, one 
character of this mineral dominates all the others: the variations in 


114 Annals of the South African Museum. 


size and in the degree of frequency of the quartz grains are independent 
of the depth at which the nodules were obtained. 

As to the “silicates” mentioned by J. Murray and A. F. Renard, 
they are undoubtedly related to rather rare felspathic elements, and 
comprise triclinic felspars. 

Glauconite.—It must be confessed that the numerous works devoted 
to this mineral and its history are far from exhausting the subject, 
as the following data show:— 

J. Murray and A. F. Renard have shown, in their description of 
the Agulhas Bank phosphates, that the glauconite, which has a 
cryptocrystalline structure like that of the glauconite of older 
deposits, is present in rounded grains independent of organisms, 
and as a product of the material filling the interior of numerous 
Foraminifera, in which it is very common. Their study has been very 
usefully completed by Messrs. L. W. Collet and G. W. Lee,* who 
discovered in these same phosphates the variety which I formerly 
designated tf as pigmentary glauconite. Bearing in mind their 
observations, I shall abstain, as far as possible, from repeating what 
is already known in this matter. 

An analysis by Giimbel { of glauconite from the Agulhas Bank 
gave the following results :— 


S10, : : . 46-90 per cent. 
Al,O; : 2. «a OGL eae 
FeO, ; + © 20-09" Shee 
FeO f : ; 3°60 y 
CaO : “20:20 
MgO . 0:70 i 
K,O : : J Os 167 a 
Na,O : EDS ee 
H,O oe eee 
99:24 wats 


In preparing for examination nodules from the Agulhas Bank, I 
have been able to gather new data, some of which concern the glau- 


* L. W. Collet, ““Les concrétions phosphatées de PAgulhas-Bank avec une 
note sur la glauconie qu’elles contiennent par G. W. Lee,” Proc. Roy. Soc. Edin., 
vol. xxv, 1904-1905, pp. 885-893; L. W. Collet and G. W. Lee, “Recherches 
sur la glauconie,”’ Proc. Roy. Soc. Edin., vol. xxvi, 1906, pp. 266-267. 

+ L. Cayeux, “Contribution a l’étude micrographique des terrains sédimentaires,”’ 
Mém. Soc. Géol. Nord, t. iv, 2, 1897, pp. 165 and 175. 

t V. Giimbel, “‘Ueber die Natur und Bildungsweise des Glaukonits,” Sitz. d. 
Math. Phys. Clas. d. k. Akad. Wiss., Miinchen, t. xxvi, 1896, p. 545. 


The Phosphatic Nodules of the Agulhas Bank. 115 


conite of the sediments whose age will be discussed later, and others 
the history of glauconite in general. 

(A) Despite the frequent occurrence of glauconite grains on the 
one hand and of Rhizopods on the other, it has not been possible to 
find a single glauconite element enclosing a Foraminifer. 

(B) The relations between glauconite and Foraminifera give 
material for interesting observations. In general, the mineral 
invades all the chambers, but it may avoid one or two, and it is also 
capable of associating there with phosphate of lime. The chambers 
once being filled, the glauconitic complex which results either ceases to 
develop—a state exhibited by absolutely intact shells of Foraminifera 
filled with glauconite—or the glauconite continues to form. If the 
latter, then a whole series of results is presented: 

(a) Exceptionally the material enters the pores, without any trace 
of metasomatism. 

(b) The glauconite fixes itself in the test, no matter where, in the 
form of minute globules, free or coalesced, which destroy the micro- 
structure of the shell. 

(c) The whole test becomes metasomatised by the glauconite, 
darker than that of the cells, which admirably preserves the structure, 
for, curious to state, the pores are not penetrated by it. Globigerinae, 
fossilised in this manner, yield beautiful tangential sections in the form 
of a network with very regular mesh, whose distinctness recalls 
sections of crinoids mineralised by ferruginous compounds. 

To these examples of pseudomorphosis of foraminiferal shells by 
glauconite—which, it must be added, are by no means rare—I add 
fragments of foraminiferal tests reduced to simple isolated arcs 
grouped in such a manner that their fragmentation was certainly 
anterior to their deposition. These too are metasomatised without 
an infilling of the pores, and yet do not show the slightest impression 
of glauconite on the matrix. In certain sections, these fragments of 
shells converted into glauconite are frequent. 

This is the first occasion on which I have been able to observe 
such a disposition to metasomatism on the part of foraminiferal 
remains without a resultant destruction of their individuality and of 
their microstructure. 

(d) In a variant of this, the whole test is metasomatised and the 
pores are invaded. Thanks to its darker colour, the surrounding 
material is perfectly differentiated from that within the shell. As 
in the preceding case, it is impossible to notice the slightest projec- 
tion into the matrix, which could lead one to suppose the existence of 


116 Annals of the South African Museum. 


a passage between the glauconitic moulds and the grains independent 
of the Foraminifera. 

(ce) In a fifth mode of occurrence—which, as in the preceding, 
involves a substitution of glauconite for carbonate of lime in Rhizopod 
shells—these appear as moulds, without trace of test. But in many 
cases there can be seen a thin border, of a very pale yellowish-green 
colour, which, between crossed nicols, is distinguished by brighter 
tints than the rest. Actually, there has been metasomatism of the 
test, and orientation of the glauconite in such a manner as to differ- 
entiate it more or less in white hght and between crossed nicols. 

This example shows that it may be necessary to discriminate casts 
of Foraminifera which may simulate whole Foraminifera, test included. 
The absence of such differentiation, which was only observed in a few 
casts, leads one to assume the existence of elements which originated 
within the cells of the Foraminifera, and were subsequently extracted 
from them by a process of mechanical decortication. This is wholly 
uncertain, and it is possible that the whole question of the inter- 
pretation of the glauconitic casts of Foraminifera may be studied 
anew. 

(C) Foraminifera, which are often profusely distributed in the 
nodules, are not the only organisms more or less altered into glau- 
conite. Leaving aside the Foraminifera, which play the principal role 
in the fixation of the mineral by organisms, we can pass at once to the 
groups of Brachiopods, Bryozoa, and Mollusca, not pausing at groups 
which are without interest. In the Bryozoa, glauconite plays a part 
in the infilling of the chambers; but most of the specimens do not 
contain it. Some Brachiopod fragments are penetrated by glau- 
conite in the form of irregular globules, which may be free, in groups, 
or fused together. 

(D) Apart from the glauconite grains associated with organic 
remains, the mineral also acts as a cement. Of this type of occur- 
rence one example only can be cited, which was observed in the most 
quartzitic nodule of all. In it was found a small stony layer of 
quartz grains accompanied by felspars, all bound together by a very 
pure glauconite which encroached on the neighbouring calcareo- 
phosphatic cement, both as irregular prolongations and as small 
completely isolated particles. 

Considering the total amount of glauconite of altered appearance, 
it can be seen that the percentage of this mineral which acts as 
gangue material reaches a figure that is wholly unknown in the 
older rocks. Those nodules which are of a brown colour and usually 


The Phosphatic Nodules of the Agulhas Bank. Ly 


angular shape possess a brown cement of various shades in which 
the obvious calcareous islets form but a very small proportion of the 
whole. 

(EK) Another variety shows glauconite present in veinlets of very 
small size, similar to those that have been noted in nodules of Albian 
age.* It will be seen later that, under certain conditions, glauconite 
in this form may have a wide distribution. 

(F) Not one of the least interesting facts to record is the profoundly 
altered appearance of glauconite in certain nodules. All things 
considered, this alteration—real or apparent—falls into two cate- 
gories which appear to denote two distinct phenomena: 

(a) In the midst of a layer rich in grains of very pure glauconite, 
one may see, é.g., a single and unique grain which is brown at the 
periphery, or several grains laden with oxide of iron to varying 
depths, the alteration sometimes reaching the centre and leaving 
only insignificant traces of the original mineral. Decomposition may 
affect all, or nearly all, the glauconite grains in a layer, thus freeing 
oxide of iron which partly remains in place and partly masks the 
neighbouring cement. The same phenomenon is produced at the 
expense of glauconite included in the organisms. In both cases, the 
iron oxide invades the gangue material and produces within it tints 
of very variable colour, a proof that the decomposition has taken 
place on sztu. 

(6) This occurrence of glauconite carrying iron oxide, playing the 
role of cement, in angular brown-coloured nodules, is probably . 
connected with another phenomenon. Nota single layer has escaped 
what appears to be, at first sight, a very characteristic alteration 
which is developed in varying degree. Some parts, which remain 
greenish, are partly masked by a little oxide of iron; others have 
become deep brown, but the colour retains a greenish shade, without 
the cryptocrystalline structure, so characteristic of the mineral, 
being destroyed. Gradually the colour turns to dark brown. All 
this takes place, therefore, as if there were a production, on an un- 
paralleled scale, of glauconite playing the part of cement, followed 
by a no less general alteration of the mineral, liberating oxide of 
iron. The phenomenon should be important, since L. W. Collet has 
found up to 21-09 and 23-70 per cent. of FeO; in the yellow nodules. f 
That is probably an incomplete picture of the sequence of events 
leading to the present condition of things. 


* L. Cayeux, op. cit., p. 181. 
+ L. W. Collet, op. cit., pp. 869 and 870. 


118 Annals of the South African Museum. 


This opinion is founded on the existence in the apparently decom- 
posed glauconitic gangue of grains of absolutely unaltered glauconite. 
On an hypothesis of one alteration, it is necessary to suppose that 
glauconite of the second generation is deeply decomposed, whilst 
that of the first generation, corresponding to the grains, has re- 
mained unaltered in all the samples examined. The conclusion, 
therefore, seems inevitable that the glauconite of the second genera- 
tion has kept its present nature right from the time of its formation. 
In other words, for unknown reasons, certain reactions caused the 
mineral to produce an excess of oxide of iron, and the so-called altered 
glauconite was in reality, at the time of its formation, nothing but 
glauconite coloured by oxide of iron. 

This question was raised by L. W. Collet, who wrote concerning 
the yellow nodules of the station Vasco da Gama: “Une idée qui 
vient naturellement 4 l’esprit est que cette couleur des nodules jaunes 
pourrait étre due a de l’oxyde de fer provenant d’une décomposition 
de la glauconie, ou que les conditions n’étant pas satisfaites pour la 
formation de la glauconie, ou hydrosilicate de fer et de potasse, il se 
soit formé un hydrate ou un oxyde de fer.”” * Accepting the fact that 
a web of typical glauconite is found behind the veil of iron oxide, as 
is shown by examination between crossed nicols, it appears more 
logical to admit that the reactions produced glauconite surcharged 
with peroxide of iron. It is not impossible that this problem likewise 
affects certain glauconites of the older deposits that appear to have 
suffered general alteration. 

Whatever the solution of this problem may be, the development 
on a large scale of glauconite as a cement-mineral is certain. 

Conclusions.—As will shortly be learned, the details enumerated 
by no means exhaust all the facies adopted by this mineral in the 
nodules. However, three salient facts have been ascertained up to 
the present: 

(A) The mineralisation of foraminiferal shells by glauconite is a 
very rare phenomenon. The specimens analysed here in detail have 
furnished examples which are incomparably more numerous than those 
of the sedimentary formations that hitherto have been submitted to 
micrographic study. 

(B) In the light of what the nodules examined have shown, no 
proof is given that the free grains originate from glauconite primi- 
tively englobed in the chambers of Foraminifera. Of the authors 
who have previously occupied themselves with the problem of glau- 

* L. W. Collet, op. cit., pp. 874 and 876. 


- 


|= 


The .Phosphatic Nodules of the Agulhas Bank. 119 


conite origin, some conclude that all the grains, of whatever kind, 
have originated in the interiors of organic cavities and, particularly, 
in foraminiferal chambers. This is, in particular, the opinion of 
Ehrenberg, W. J. Sollas, Bonney, J. Murray, and A. F. Renard, etc. 
Others, such as J. Bailey, A. E. Reuss, V. Giimbel, etc., believe in a 
dual mode of formation. 

The hypothesis—often propounded—that the innumerable elements 
whose shape does not suggest organic origin result from the irregular 
growth or fusion of casts of Foraminifera receives no support, in 
spite of the fact that the occurrence yields observations particularly 
decisive in this respect because of the large number of nodules and 
of free grains found side by side. No example of a cast passing into 
an irregular grain has been seen, and no example of an irregular 
grain enclosing a foraminifer with its chambers obliterated by 
glauconite. In every case, without exception, the two types of 
glauconite grains present are entirely independent; my absolute 
conviction is that, in this deposit, glauconitic casts remain glauconitic 
casts, and that grains, whatever their size and form, never originate 
from them. Many examples, taken from elsewhere, help to give this 
conclusion a general application. I add that the development of the 
epigenic glauconite of the cement and a mode of occurrence on which 
I shall lay stress later prove, once more, that glauconite can be formed 
on a large scale independently of organisms. 

(C) Incomplete as it is, the preceding study shows the existence of 
at least two generations of glauconite—one consisting of the free grains 
and casts of Foraminifera, and the other the epigenetic glauconite of 
the matrix and the rare veinlets. 


2. Organisms.—Beginning with the highest forms, it is necessary 
only to mention the exceptional presence of splinters of bony tissue, 
which are usually absent. 

To the Brachiopoda are assigned a small number of fragments of 
tests and of entire shells which, on the average, number at the most 
one example in each section. Both have preserved their structure 
intact, and the canals may, or may not, be filled with phosphate of 
lime. 

Less rare than the Brachiopods, Bryozoa—represented by very 
fragmentary colonies—do not appear in all the thin sections; their 
maximum frequency is 20 individuals in one section—a notable 
proportion, having respect to the size of the débris. 

The Mollusca, which form the most important group after the 


120 Annals of the South African Museum. 


Foraminifera, are chiefly found in the angular brown nodules. There 
occur some dozens of curved, fragmentary, and usually thin forms 
with their microstructure preserved. 

To the Echinoids are assigned some calcareous plates, with a single 
optical orientation, whose microstructure is almost entirely destroyed, 
as well as some exceedingly rare spines. All occur in the brown 
nodules, to the maximum number of a dozen plates in a section. 

The very rare Sponge spicules observed are calcified. The one 
example, unique up to the present time, proves that the solution of 
siliceous spicules and the replacement by calcite are not necessarily late 
phenomena posterior to the emersion of the sediments. 

The Foraminifera, however, easily surpass the preceding organisms 
in importance and numbers. Except in the brown nodules with 
Bryozoa and large benthic Foraminifera, Globigerinae predominate, 
to the point of forming by themselves the whole Rhizopod fauna. 
Generally speaking, the Foraminifera are whole and beautifully pre- 
served. With the exception of the brown nodules, rich in bottom- 
dwelling Foraminifera, the shells, both entire and fragmentary, show 
no corrosion, and it is impossible to imagine the fragmentation of the 
Globigerinae otherwise than by mechanical action. 

Boring organisms have left profound traces in the brown nodules 
in the form of numerous irregular tubes filled with phosphate of 
lime, ferruginous glauconite, or opaque oxide of iron, most of which 
are probably attributable to sponges. -Except for the sponge 
spicules and the Echinoid remains, all the groups have suffered from 
their action. 


3. Cement.—From the point of view of the constitution of the 
matrix, the nodules form very dissimilar groups—the brown nodules 
on the one hand, and the blackish nodules on the other. 

(A) By their matrix the latter recall immediately, under the micro- 
scope, the phosphatised chalks of the North of France. Although 
on the average little developed, on account of the profusion of 
Foraminifera, the cement is generally calcareo-phosphatic, at places 
phosphatic and calcareous and, very exceptionally, formed of pure 
phosphate of lime in very restricted areas. In white light, the 
phosphate of lime is identified by a very pale yellowish tint, which 
becomes most pronounced as the phosphate content increases and, 
finally, attains a pale yellow colour. Under crossed ricols, as a rule 
the yellowish base becomes charged with very numerous small brillant 
and iridescent spots, representing the presence of carbonate of lime, 


The Phosphatic Nodules of the Agulhas Bank. 121 


which are prominent when phosphatisation is feeble but disappear 
completely in the exceptional layers which are wholly phosphatic. 
In short, it is necessary to examine but a very limited number of 
nodules to observe a complete passage from a matrix exclusively 
calcareous to a cement which is entirely phosphatic. It follows 
that, on the average, these nodules are rather poor in phosphoric 
acid. 

(B) Actually, the brownish nodules—which differ considerably 
from the preceding—are far from forming a constant variety. 
Although limited to two specimens, my studies have shown profound 
differences between them. 

The one which is considered a partly phosphatised Globigerina ooze 
possesses a cement that is essentially glauconitic and more or less 
haematitised (Pl. XX XIV, fig. 3), enclosing some plates of Globigerina 
mud free from mineralisation, exceptionally converted into mostly 
crystalline calcite, and others—rather rare—which are on the way to 
being phosphatised. It is difficult to say if the matrix, which is 
strongly impregnated with oxide of iron, is not at the same time 
slightly phosphatised, since the brown base prevents the identifica- 
tion of phosphate of lime. 

The cement of those brownish nodules that are rich in large benthic 
Foraminifera has visible phosphate of lime in preponderance. Some- 
times this mineral surrounds uniformly all the perfectly preserved 
constituents in a yellowish base, which is undifferentiated and full 
of minute calcareous specks; sometimes it is associated with calcite, 
which is mostly crystalline, and forms a thin, pure yellow border to 
all the material which it serves to bind. In this variety the organic 
remains are corroded and are often almost unrecognisable. Itis rather 
the calcite which actually plays the part of cement. 

The same slide shows both these modes of occurrence, to which can 
be added a third, characterised by a ferruginous matrix (Pl. XX XIV, 
fig. 4) like that of the preceding specimen, which does not permit 
one to say that the oxide of iron is intimately associated with glau- 
conite, although this is probable. This ferruginous matrix may also 
enclose calcite in the form of irregular patches of varying size. 

The history of these nodules is thus complicated by a profound 
alteration of the original constituents of the ooze which have escaped 
phosphatisation and mineralisation, these constituents having now a 
crystalline condition comparable with that of the highly recrystallised 
ancient limestones. There are even true veinlets of calcite in the mass, 
like those found in ancient marbles. These are, it is true, few in 


122 Annals of the South African Museum. 


number, and one cannot invoke dynamic alteration to account for 
them. 

Mode of Occurrence of Phosphate of Lime.—At the outset, phosphate 
of lime plays a double réle: 

(A) It fills the chambers of Foraminifera, the cellular fabric of 
echinoderm plates, the cells of Bryozoans, etc. Actually, this mode 
of occurrence 1s bound up with the following, for the phosphate of 
lime in the foraminiferal chambers retains traces of calcite which 
bear witness to epigenetic phenomena. The truth is that, despite 
appearances, phosphate of lime is not concentrated as such in these 
spaces. 

(B) It is produced on a large scale by alteration of the calcareous 
matrix of the foraminiferal muds and, on a reduced scale, in the 
tests of the organisms present. 

It follows that, in its two principal modes of occurrence, phosphate 
of lime is undoubtedly a substitution product. 

The phosphate of lime formed at the expense of the cementing 
material is optically amorphous and lacks any kind of morphological 
differentiation; it therefore belongs to the group of colophanite or, 
more exactly, of fluocolophanite. This is so for all the nodules ex- 
amined to date. It will be shown later, from a study of aberrant 
specimens, that a crystalline variety may also occur, but seemingly 
exceptionally. 

Granular phosphate of lime is extremely rare, when it is not 
completely absent. There are doubtless exceptions to this rule, for 
J. Murray, examining material dredged by the Blake, noted that a 
concretion, obtained in the Atlantic at Station 317, in lat. 31° 57’ S., 
contained phosphatic grains “similar to the grains found in the 
phosphate nodules dredged near the Cape of Good Hope and elsewhere 
by the Challenger.” * 

Finally, the alterations undergone by the parent rocks, converted 
into poor phosphates, are much less profound than those noted by 
J. Murray and A. F. Renard in the nodules from the Indian Ocean 
where, at the same time, are integral substitution, destruction of 
organic characters, development of zonary structure, and crystallisa- 
tion of the phosphatic material. 


* J. Murray, “Report on the Specimens of Bottom Deposits,” Bull. Mus. Comp. 
Zool., vol. xii, 1885-86, p. 43. 
+ J. Murray and A. F. Renard, op. cit., pl. xx, figs. 3 and 4. 


The Phosphatic Nodules of the Agulhas Bank. 123 


ANOMALIES OF COMPOSITION AND STRUCTURE OF 
. NODULES. 


Among the nodules of dark colour which arise from quartzitic 
Globigerina muds are some that are characterised by the presence of 
inclusions, traces, and incrustations of aberrant constitution, and some 
by the juxtaposition of two very dissimilar rocks. 

1. Nodules with very Glauconitic Inclusions.—(A) Glauconite 
gives rise to narrow bands with well-defined contours, from which 
minerals and organisms are absent. These differ from the grains 
disseminated through the rock by a light brownish colour, indicative 
of an elimination of peroxide of iron. In this form the mineral 
presents its true appearance. 

This same variety serves as a cement to large grains of glauconite 
of a normal colour, strikingly different from those that form an in- 
tegral part of the nodules. Thus it is possible to see well-defined bands 
formed exclusively of glauconite grains of one shade (g), surrounded 
by a glauconitic matrix of an entirely different shade (Pl. XXXV, 
fig. 1). Thus there are present two generations of glauconite which 
have nothing in common, in point of age, with those concerning 
which question was raised in the study of nodules of normal type. 
These can, moreover, be seen in a given nodule. 

(B) The inclusions are rich in grains of ordinary glauconite, which 
may form the greater part of them, and often touch one another. 
Apart from insignificant exceptions, the mineral is very clear, except 
when it is obscured by the neighbouring plates. 

With few exceptions, the inclusions in this category show no 
trace of Foraminifera or other organisms. Their cement is entirely, 
or almost entirely, phosphatised and cryptocrystalline, i.e. it is a 
staffelite. In general, the phosphate is differentiated as a very clear 
border round grains of glauconite and of quartz. This differentiation 
is also observed, but not always, around the islets and bands. 

The relations of these inclusions to the neighbouring rock are 
illustrated by the two following examples :— 

In the first, there is a beautifully distinct band of glauconitic 
phosphate which is bounded on one side by a curved line traversing 
a large plate without deviation, cutting as with a knife the glauconite 
grains and phosphatic constituents which it encounters. On the 
other side, the boundary is irregular and roughened with protuber- 
ances and hollows in a manner that is immediately reminiscent of the 
phenomena of corrosion. Moreover, along this line of separation the 


ia | 


124 Annals of the South African Museum. 


nodule is strongly impregnated with glauconite, just as is frequently 
found on old hard and corroded floors. 

But the relations between these two portions are susceptible of 
quite another interpretation. The cementing of the two rocks follows, 
then, a very irregular line, whose limits it is often impossible to fix, 
and the rocks merge into each other at their junction. 

From the study of a number of thin sections it is possible to obtain 
an idea of the form of this sort of inclusion. They represent, in fact, a 
product of the partial or complete infilling of cavernous or scoriaceous 
holes in the nodules. Such of these bands as can be followed through- 
out their whole development leave the bottom of a hollow in the form 
of a funnel and penetrate into the heart of the nodules. In short, 
bands and islets are really the same, and simply represent respectively 
longitudinal and transverse sections of cavities filled by greensands. 

The specimens which favour this type of observation correspond, 
I believe, to the concretions “avec fentes remplis d’une substance 
verdatre ’’? which appears to L. W. Collet “pouvoir étre attribuée a 
de la glauconie.”’ *- 

Nothing is more evident than that this material exhibits an inter- 
penetration of two fundamentally distinct types of rock—an older, not 
typical, Globigerina mud and a true greensand, both consolidated. 
The same contrast exists between the material of the nodules and their 
mineral incrustations, which do not differ at all from the islets and 
bands. 

Under these conditions it 1s impossible to escape from the con- 
clusion that the nodules, formed at the expense of non-typical Globigerina 
muds, have been removed from the locality in which they were formed and 
deposited in areas where greensands were being laid down. From this 
conclusion is derived another: In the process, the nodules have been 
transported from one sediment more pelagic than terrigenous into another 
plainly terrigenous, and from greater depths to more shallow areas. 


2. Nodule formed of two differing Rocks (Pl. XX XIII, fig. 1, and 
Pl. XX XV, fig. 2).—Station 10 (Vasco da Gama), situated 25 km. from 
the coast, has yielded a set of “‘concretions,”’ among which is the largest 
known, as well as rolled pebbles of quartzites and schists. In one of 
the nodules which he has figured, L. W. Collet has clearly distinguished 
two portions separated by a black line 0-5 mm. thick, one of which 
is yellowish and characterised by numerous Foraminifera, the other 
blackish and rich in grains of glauconite.t This nodule, whose 


* L. W. Collet, op. cit., p. 867. 
+ L. W. Collet, op. cit., p. 869 and fig. 3. 


The Phosphatic Nodules of the Agulhas Bank. 125 


greatest diameter is a dozen centimetres, is formed of two distinctly 
different rocks, joined in such a manner that it would not be correct 
to say that one serves as a matrix for the other. 

On a polished surface (Pl. XX XIII, fig. 1) there can be seen a 
lower part of clear yellowish colour, darkened above over a width of 
about 1 cm., and an upper part of very deep tint in which, under the 
lens, a crowd of little black glauconitic grains can be distinguished. 

(A) The yellowish portion (Pl. XX XV, fig. 2, a), which was clearly 
the lower part of the specimen in place and which, according to L. W. 
Collet, was “‘formée presque entiérement de coquilles de Foramini- 
féres,’’ must have a variable composition. Actually, sections cut from 
the same specimen display but a small proportion of Foraminzfera 
and, it is interesting to note, an irregular distribution of the organic 
constituents—a fact which doubtless explains the discrepancies 
between our observations as to the quantity of Foraminifera present. 
However, as far as I have seen it, the yellowish rock contains a pre- 
ponderance of fragments of Bryozoa and, in addition, Foraminifera 
which are rare or relatively frequent and mostly Globigerinae. 
Echinoderms are represented by several plates and one or two echinoid 
spines, together with a single holothurian spicule. In addition there 
are some indeterminable fragments. Along certain widespread planes 
all the organisms are represented by hollow moulds. 

~The matrix of the rock, which in each section is formed of lime- 
stone containing dozens of quartz granules, is more or less pene- 
trated by phosphate of lime and glauconite; it becomes more phos- 
phatic as one approaches the contact of the two rocks. At the point 
(b) and along the suture-line (c) it is wholly phosphate and glauconite. 

The phosphate of lime, tinted in yellow of differing shades, is 
clearly cryptocrystalline. In the sections showing the onset of de- 
calcification it is crystalline in the form of short scales bordering the 
cavities or arranged in very thin zones. Hxcept for a short distance 
from the line of contact (c) carbonate of lime is abundant, even in the 
most phosphatic portions, in the form of a very large number of 
small corroded fragments which are identifiable. The phenomenon 
of epigenesis is clearly marked. 

Everywhere glauconite accompanies the phosphate of lime as very 
scattered grains, and chiefly as pigment. Its partial alteration is con- 
comitant with the yellow coloration of the deposit. The fact that this 


does not extend to the glauconite of the upper part of the nodule, 


which is of more recent formation, shows that the decomposition of 
this mineral dates from far back and that it is of submarine origin. 
VOL. XXXI, PART I. «414 


od 


126 - Annals of the South African Museum. 


The dark zone (c), which marks the upper limit of the rock con- 
taining Bryozoa and Foraminifera, is characterised not only by the 
complete disappearance of carbonate of lime, but also by a develop- 
ment of pigmented glauconite on a scale which is unique in my experi- 
ence. The phosphate of lime and glauconite are intimately associated 
there. The latter mineral is concentrated in plates, pockets, and 
bands that are arranged very irregularly. Some clear areas are a 
kind of discontinuous network formed of anastomosing veinlets, 
and have no parallel in any known rock. In addition, glauconite 
metasomatises all the organisms in the zone, such as Foraminifera, 
Bryozoa, and Echinoderms, showing absolute freshness everywhere. 
It is here that the most beautiful examples of Foraminifera converted 
into glauconite can be seen. 

The same line of suture (c) can be clearly traced from one end to the 
other. It is characterised sometimes by a concentration of coloured 
glauconite whose density may or may not decrease lower down, and 
sometimes by a concentration of phosphate of lime marked by a 
particular coloration and transparency. There is no encroachment of 
phosphatised limestone beyond this line. Finally, its shape is marked 
by a number of irregularities which give it the characteristic shape of 
a line of corrosion. 

(B) Beyond this line an entirely different rock appears. This is 
coarse-grained, crowded with glauconite, much more quartzitic, 
lacking invertebrate remains, and has a phosphate matrix. The 
contrast is extremely striking. Here is no trace of pigmentary 
glauconite or of epigenesis. The mineral forms a crowd of very 
pure grains of the ordinary type, measuring on the average 1 mm. 
in diameter, and may preponderate in a good many of the sections 
(Pl. XX XV, figs. 2 and 3). In all characters this glauconite is identical 
with that of the incrustations and of the nodules previously described. 
In addition to the granules which represent quartz in the preceding rock, 
there are here large grains measuring 0-6 mm. and more (Pl. XX XV, 
figs. 2 and 3, q), of such a kind that, in one and the same section, the 
diameter of the grains may vary in the ratio of 1: 30 and even more. 
This is quite abnormal and necessitates a strongly agitated environ- 
ment. Organisms are represented only by one or two fragments of 
bony tissue, with very undulose extinction. The matrix, composed 
of phosphate of lime which is grey, pale yellow, or deep yellow, is 
isotropic, or cryptocrystalline, between crossed nicols and nearly 
always free from fine calcareous inclusions. In its characters this 
rock might be taken for certain pebbles of the greensands with 
Douvilleiceras mammillatum from the east of the Paris Basin. 


The.Phosphatic Nodules of the Agulhas Bank. 127 


It can be noted here that this constitution is a replica of that of the 
nodules formed exclusively of phosphatised greensand, described with 
the six types already analysed (p. 112). 

This specimen yields good evidence of a great disturbance which 
has affected the sea-bottom at some time and produced an eroded 
surface. There are two fundamentally different rocks in contact, 
one arising from greensands and the other from a bryozoan fora- 
miniferal limestone, two deposits formed in different surroundings. 
In parenthesis, the term “concretion” does not appear the correct 
one to designate a rock detached from the junction of two different 
- formations and shaped into a nodule or, more properly, a pebble. 

Corrosion of Nodules.—The intrusion of glauconite of the greensands 
into certain nodules, discussed above, has arisen from true corrosion 
phenomena which, as a final product, have resulted in the formation 
of scoriaceous material. The cavities thus created are to-day either 
empty or have been more or less filled with glauconitic phosphate. 

The material at my disposal is of too restricted a nature to elucidate 
the problem of the corrosion of pebbles. Some nodules have escaped 
corrosion, but the proportion is unknown; this is the case for those 
which have been produced from greensand. Some have been affected 
over their whole surface, resulting in a scoriaceous appearance. 
Others have been worn on one side only; e.g. a brownish, very angular 
nodule resting on the bottom on a flat face has been corroded and 
finely perforated on its upper side only. The conclusion is reached 
that corrosion has affected material that was already detached from 
the parent rock, 7.e. the nodules themselves, in which the phenomenon 
has nothing in common with the grooving shown by one specimen. 

It cannot be questioned that all the corrosion is anterior to the 
penetration of the greensands into the cavities created by it; 
but it is no less certain that the wearing away was continued 
after the change in environment which caused the greensands to 
appear. As witness to this, there is the nodule formed of two distinct 
rock-types, of which the upper part, characterised by an old phos- 
phatic greensand (Pl. XX XIII, fig. 1), is itself corroded. 

At the moment this phenomenon of submarine corrosion requires 
elucidation, especially as to the nature of the environmental conditions 
that are favourable to it. Possibly an examination of a large number 
of specimens would throw some light upon it by supplying an answer 
to one important question—Does a single station provide, at one and 
the same time, corroded pebbles and pebbles that have escaped 
corrosion ? 


128 Annals of the South African Museum. 


Nomenclature of the Phosphatic Material of the Agulhas Bank.—1 
have purposely refrained from using the term “concretion,” since 
it was first necessary to obtain discriminatory knowledge of the 
mode of origin of the material. In its general acceptation the term 
nodule does not imply any given mode of origin, while that of 
concretion immediately suggests concentration around one or several 
points of attraction. Such is the idea that is held—often wrongly— 
of the mode of formation of flint. Among this material, the only 
true concretions are the phosphatic greensand nodules. All the 
rest are referred to a phenomenon of substitution of phosphate of 
lime for carbonate of lime, a phenomenon which does not obey the 
same laws as the former and which, moreover, is quite independent 
of their morphology. 

In short, all the phosphatic materials of the Agulhas Bank, without 
exception, are nodules, and an unknown fraction of these nodules falls 
within the category of concretions. 


INTERPRETATION OF THE FACTS, AND CONCLUSIONS. 


In describing the material dredged to the south of the Cape of Good 
Hope which furnished almost the only basis for their study of the 
mode of formation of phosphatic concretions, J. Murray and A. F. 
Renard laid emphasis on the important fact that the minerals and 
organisms of the nodules on the one hand, and of the sediments in 
which they are included on the other hand, are identical, both in the 
case of the concretions of the greensands and those of the Globigerina 
mud. This is to say that all the present-day phosphates, and those 
of South Africa in particular, have been formed in situ. In this 
connection there is no doubt that J. Murray and A. F. Renard 
studied material, formed in place, that perfectly displayed the 
characters of the greensands and of the Globigerina ooze. 

L. W. Collet agreed with their opinion, writing ‘‘ Les concrétions 
phosphatées sont en quelque sorte ’image du fond dans le quel on les 
trouve.’ * There are certain facts cited by L. W. Collet in his deserip- 
tion of the phosphatic concretions of the Agulhas Bank which support 
this contention. One is the observation that in the concretions “la 
quantité de coquilles de Globigérines augmente avec la profondeur de la 
mer.” 

It is pertinent here to state that the concretions studied by J. Murray 


* BE. W. Collet, Les dépots marins, 1908, p. 200. 
+ L. W. Collet, Proc. Roy. Soc. Edin., vol. xxv, 1905, p. 879. 


| = 


The Phosphatic Nodules of the Agulhas Bank. 129 


and A. F. Renard are not the same as the large number of specimens 
obtained by the Cape Government. Having stated this, it appears 
to me that a conclusion contrary to that given above is supported 
by some very convincing evidence and by arguments drawn from the 
interesting monograph of L. W. Collet on the phosphatic concretions 
of the Agulhas Bank. 

Station 12 (421 m.) of the Agulhas Bank yielded L. W. Collet some 
nodules rich in Globigerinae, which were frequently whole, and some 
which “se composent uniquement de grains de glauconie cimentés par 
le phosphate.” * Similarly, Station 7 (146-238 m.) } yielded some very 
ferruginous nodules of a yellow-brown colour and others “‘ressemblant 
extérieurement 4 de la ponce.” { The first, according to G. W. Lee, 

- are free from detritic minerals, and the second contain abundant 
quartz, etc. Lee concludes, logically enough, that “il y a une 
différence essentielle entre ces deux espéces de concrétions.” § 

My own observations show that the brownish nodules, dredged at 
one and the same place, are of very differing characters (p. 112); 
one type is a consolidated globigerinal mud, the other is formed 
from a deposit of large benthic Foraminifera with fragments of 
bryozoan colonies. These two very different rocks cannot have been 
formed at the same depth. 

The existence of cavernous nodules with Globigerinae carrying in- 
clusions from which these tests are excluded, as well as that of the 
nodule from Station 10 formed of two distinct rock-types, are further 
evidence on the same lines. 

From this assemblage of facts the following conclusions can be 
drawn :— 

(1) All the nodules used in this study and, without doubt, a portion 
of those studied by L. W. Collet have, at some period of their history, 
become remanié. We shall see later how it is possible to reconcile 
the presence of nodules formed on the spot with the existence of 
transported pebbles. 

(2) To this idea of submarine remaniement can be added another, 
also formulated for the first time, which concerns the present-day 
sea-bottom and can be called a submarine erosion. The composite 
nodule, described above, furnishes proof that the bottom of the sea 
has suffered denudation, and that this erosive action coincided with 


* L. W. Collet, op. cit., p. 876. 
j{ This Station 7 is not the same as that given in the table on p. 106. 
t L. W. Collet, op. cit., p. 877. 
§ L. W. Collet, op. cit., p. 878. 


130 Annals of the South African Museum. 


a radical change in the nature of the surroundings as well as in that 
of the sediments. The phenomena of erosion have acted on the cal- 
careous substratum in the same way as the changes of level of the supra- 
Cretaceous epoch on the chalky muds of the Paris Basin ; hardening 
of the bottom, impregnation by phosphate and by glauconite, etc., are 
present in both cases. 

It remains to determine the succession of events that can explain 
this deposit, a problem that the preceding discussion seems to have 
obscured, and which necessitates a departure from the domain of facts 
for a moment. 

In my opinion, the problem is essentially dependent on the following 
important fact: All the remanié material is derived from deposits that 
were formed at a greater depth than that at which they were dredged. - 
This fundamental observation leads to the conclusion that the region 
in question has been the seat of important bathymetrical changes. 
Without justifying their opinion, J. Murray and E. Philippi have ad- 
vanced the idea of an elevation of the sea-floor.* Unless I am in error, 
everything points to an uplift of great amplitude, which has raised up 
lumestones of varied types, and even Globigerina oozes. This change of 
level, which is to be measured in hundreds of metres, has terminated 
the deposition of limestones, has inaugurated the formation of green- 
sands, and has induced the phenomena of erosion, of disintegration, 
transportation, and re-cementation of the material. Phosphatisation 
enters into this complex of phenomena, of which we find an equivalent 
in the distant past, always in close connection with the manifestation 
of great changes. It is, too, from this period that the phenomenon 
of corrosion, which has shaped a portion of the nodules, dates. 

To be more precise, it must be added that the phenomenon is less 
simple than it appears, for it is complicated both by a delay in 
movement of the material and by a resumption or continuance of the 
corrosive action after the change in level, as is shown by the nodule of 
mixed composition. 

Nothing is easier to understand than that this change of level, 
with its consequences, can explain the duality of origin of phosphatic 
rocks. At the same time as the pre-existing materials have been 
metasomatised there were formed true concretions at the expense 
of the greensands that were in course of formation. I shall show 
later, in examining the more ancient phosphates, that such an 
association is not unique in the sedimentary series. 

Although I believe this to be the true succession of events, it must 


* J. Murray and E. Philippi, op. cit., p. 186. 


The Phosphatic Nodules of the Agulhas Bank. 131 


be confessed that it is not wholly clear. One episode of the history 
in particular remains a puzzle. How did the nodules, or the limestone 
fragments from which they were formed, become concentrated in 
one area? This question is all the more important, seeing that rocks 
formed under differing bathymetric conditions are found to-day 
gathered together in one place. 

Without pretending to give to this problem the answer it deserves, 
it may be that the solution should be sought in the following 
direction :— 

One finds, on the deep-sea side of the Agulhas Bank, very steep 
slopes which sink rapidly to depths of 3000 metres and more (see Map, 
Pl. XXXII). Actually, all the concretions dredged by the Challenger, 
and most of those supplied to L. W. Collet by the Cape Government, 
came not from the Agulhas Bank proper but from the external slopes. 
Of the nine stations which supplied the nodules sent to me one only 
is on the actual Agulhas Bank. It is from this that were obtained the 
brownish nodules, one of which corresponded to a Globigerina mud, 
and the other to a calcareous clay enclosing numerous benthic 
Foraminifera and Bryozoa. To clarify the idea even a little, it can 
be presumed that the different types of limestone from which the 
phosphatic nodules have been derived outcrop in horizontal beds of 
the slopes of the plateau which is covered with a mantle of green- 
sands, which is probably very thin. Whatever may be the succession 
of the deposits it is natural that, following the change of level, an 
agglomeration will be produced in the talus on the outside of the 
ridge. Further, it must be supposed that this surface, profoundly 
eroded during uplift, would enable the dredge to explore different 
horizons of the complex forming the Agulhas Bank. 

Whether this view is acceptable or not, it is indisputable that the 
phosphatic nodules of the Agulhas Bank are not dependent on present- 
day phenomena, a conclusion contrary to the opinions of J, Murray, 
A. F. Renard, and L. W. Collet. 

Possibly the conclusion, stated thus, is too absolute. It includes 
all the nodules which do not result from the consolidation of green- 
sands. It applies equally to the nodule formed of two rocks fused 
together, one of which was derived from phosphatised greensand. 
Actually, one can show that the nodules formed from metasomatised 
limestones are not of modern origin, but it is not possible to show 
this for the concretions that are exclusively built up in consolidated 
greensands. It is not an objection that the remaniement of the 
fragment detached from the contact of the greensands and the 


132 Annals of the South African Museum. 


underlying calcareous beds leads, naturally, to the supposition that 
those nodules which were produced solely from the greensands date 
from the same period as the others. If I do not state my conclusions 
as absolute, it is because I can picture an hypothesis which involves 
a prolonged period for the genesis of the greensand nodules. 

It is not possible to fix the age of the phosphatic materials of the 
Agulhas Bank from a knowledge of their mode of origin. In this 
connection it must be recalled that the dredgings have not furnished 
only phosphatic nodules. L. W. Collet taught that the nodules of the 
Agulhas Bank are “accompagnés de nombreux restes d’animaux,”’ 
“consistant en dents de poissons et de squales, os tympaniques de 
cetacés.”” * We know also from him that, at a depth of 421 metres 
(Station Vasco da Gama),{ there were dredged, in addition to the 
concretions, “un grand nombre de fossiles de Lamellibranches et de 
Brachiopodes, ainsi que des coquilles mortes,”’ and living representa- 
tives of several groups “qui témoignent d’une abondance de vie” ¢ 
in this region. Concerning the fossils, L. W. Collet notes that one finds 
“tous les modes de passage depuis la coquille morte, blanche, a celle 
complétement transformée en phosphate de couleur brillante brune.”’ § 
The same author records the presence of phosphate of lime in the 
dead shells. Unfortunately, none of the fossils are identifiable. 

Now that the problem has been placed on a new basis, the shells 
ought to furnish material for two interesting observations: (1) If 
they are corroded, they must date at least from the period of dis- 
turbance which determined a radical change in the nature of the 
sediments. (2) If showing no trace of corrosion, and if they are of the 
same size and thickness as the living shells or the unfossilised dead 
ones, then the hypothesis of remaniement (derivation) must be dis- 
carded, and cannot be used in fixing the age of the deposit. 

Sir John Murray and E. Philippi, whose brief account of the 
phosphates of the Agulhas Bank appeared three years after the 
publication of L. W. Collet’s memoir, wrote: “Es scheint aber sicher 
zu sein, dass diese Fauna nicht alter als jungtertiar ist. . .”’ || without 
furnishing any proof. 

The presence of a tooth of Carcharodon, probably belonging to the 

* L. W. Collet, Les dépdts marins, 1908, p. 207. 

+ This station has furnished nodules rich in Globigerinae, as well as others 
that are solely composed of grains of glauconite cemented by phosphate of 
lime. 

t L. W. Collet, Proc. Roy. Soc. Edin., vol. xxv, 1905, p. 875. 


§ L. W. Collet, zbid., p. 876. 
|| Sir John Murray and Professor E. Philippi, op. czt., p. 186. 


The Phosphatic Nodules of the Agulhas Bank. 133 


living C. Rondeletw, does not settle the question. Itis useful to note 
that Dr. A. W. Rogers, formerly Director of the Geological Survey of 
the Union, recognised among the material obtained by Dr. Gilchrist 
a shell of Aturia partially enclosed in the phosphate “which,” he 
wrote to me, “reminded me of the remaniés fossils in the Cambridge 
Greensand.” This observation would date the deposit in a singular 
manner, if the hypothesis of derivation (remaniement) could be dis- 
carded with certainty, for the most recent known forms of Aturia 
appear in the Miocene. Butif at least part of the material is derived, 
then any fixed conclusion is impossible. 

Whatever the age of the phosphatic deposits of the Agulhas Bank, 
it is indisputable that the major phosphatisation which produced 
them began before the present epoch and after the disturbance 
which put an end to the deposition of limestones and substituted 
greensands for them. The change of level is such that it is probably 
necessary to place it before the Pleistocene period. 

The fact, observed by J. W. Collet, that dead shells belonging to 
living species are encrusted with phosphate of lime does not lead to 
the conclusion that the phosphatisation has been prolonged until 
quite recent times. The absence of incrustations of phosphate on 
_ living shells rather leads to the supposition that deposition has not 
continued until now. 

Perhaps the problem of the age of the phosphatic nodules could be 
approached from an entirely different standpoint. Has the change 
in level shown by these submarine sediments, not far from the coasts, 
left traces on the dry land? If so, one can foresee the possibility of 
solving the problem. This aspect of it must be dealt with by those 
geologists who have such a wide knowledge of that portion of South 
Africa. 

From all that has been learned, it results that the submarine deposit 
of the Agulhas Bank constitutes in some way the last phase of an wmportant 
series of phosphatic deposits which are arranged almost from one end 
to the other of geologic tume. From this it follows that the deposit in 
question, that must be considered whenever one seeks to explain 
the origin of sedimentary phosphates, is far from presenting us with 
a clear picture of all the factors which play a part in their formation. 
As far as is known at present, it shows no essential difference from the 
older phosphatic formations. 

It is no exaggeration to say that the preceding study has been 
unexpectedly fruitful and full of novelty. Apart from the ideas of 
great changes of level, and of submarine erosion and derivation 


134 ._ Annals of the South African Museum. 


which have clearly emerged from it, it has shown the existence of 
several generations of glauconite and of instructive conditions of 
fossilisation (calcified Sponge spicules and numerous Foraminifera 
with glauconitic tests). Moreover, it has made known phenomena, 
developed in a submarine environment, which ordinarily belong to 
continental areas, such as decomposition of glauconite, the sometimes 
widespread crystallisation of carbonate of lime which has escaped 
phosphatisation, the genesis of true veinlets of calcite, decalcification 
observed in the composite nodule, and corrosion of numerous nodules. 
With regard to the problem of genesis, this study has stressed the 
extreme rarity or absence of microscopic remains of fishes, organisms 
whose role is usually held to be preponderant in the genesis of phos- 
phate nodules. Finally, from the chemical point of view, it has 
shown the presence of fluorine, and thus created a close connection 
between the phosphates of the Agulhas Bank and those of earlier 
times. 

Thanks to the material placed at my disposal, this description has 
assumed the character of a study of submarine geology applied, not 
to an interior sea of reduced depth like the English Channel, but 
to one of sub-oceanic depths, with both pelagic and terrigenous 
sediments. From every aspect it is intimately bound up with the 
geology of emerged formations. 

If one separates phosphatic nodules of the Agulhas Bank from 
those of the present day for the reason that they are older—with 
the reservation made above in favour of the pebbles that are solely 
derived from the greensands—the distribution of phosphates of 
modern origin becomes very restricted. Hitherto, nearly all the 
evidence of the properties of modern phosphates has been furnished 
by the nodules of the Agulhas Bank. If these latter are considered 
as of an earlier origin, all that remains within the category of phos- 
phates of modern origin are the nodules dredged by the Blake and the 
Challenger in areas not within the Agulhas Bank; and the conclusions 
reached for the Agulhas Bank nodules are of such a nature as to 
suggest doubts as to the true age of the others. 

Moreover, the authors who have recorded the existence of phos- 
phatic nodules have scarcely given all the necessary information 
required. It is impossible to find statements as to their degree of 
frequency; nor do we know whether we are dealing with petro- 
graphic curiosities or with true pebble-beds—a matter of considerable 
importance. If the former—which is more probable—the existence 
of a few nodules throws no more light on the genesis of sedimentary 


The Phosphatic Nodules of the Agulhas Bank. 135 


phosphates than does a study of the scattered nodules in the White 
Chalk of the Paris Basin clarify the problem of the origin of the older 
phosphates. 

Finally, almost nothing is known of the deposits of nodules other 
than that of the Agulhas Bank. What is known can only justify 
doubts as to their age. The single nodule, described by J. Murray, 
which was dredged by the Blake,* came from a hard bottom, 1.e. from 
unusual conditions of sedimentation. The question arises, Is not 
this hard bottom evidence of a change of level with all its conse- 
quences? That such a question can be asked is enough to show that 
the value of the nodules dredged by the Blake is not known. 

Contrary to expectation, the present-day formations throw but little 
light on the vast problem of the genesis of sedimentary phosphates. 
To find a solution it is necessary to pin our faith on the past. 


* J. Murray, Three Cruises of the Blake, vol. i, 1888, p. 276, fig. 189. 


136 Annals of the South African Museum. 


EXPLANATION OF PLATES. 


Prate XXXII. 
Map of Agulhas Bank area. 


PuaTtE XXXII. 
Morphology of nodules, nat. size. 


Fig. 1. Rounded nodule, formed by two different rocks (see Pl. XXXYV, fig. 2). 
Fig. 2. Scoriaceous nodule, Stat. 7, Cape Point, 560 fath. (1024 m.). 
Fig. 3. Very irregular nodule, Stat. 6, Cape Point, 315-400 fath. (576-732 m.). 


PLATE XXXIV. 
Thin sections of nodules. 


Fig. 1. Calc-phosphate nodule, very quartzitic. Stat. 8, Cape Point, 300 fath. 
(549 m.). gq, quartz; g, glauconite. (x 50.) 

Fig. 2. Calc-phosphate quartzitic nodule, rich in Globigerinae. Stat. 3, Vasco da 
Gama Peak, 230 fath. (421 m.). 9g, quartz; g, glauconite. (x 45.) 

Fig. 3. Phosphatised and ferruginous nodule, full of Globigerinae. Stat. 5, Cape 
St. Blaize, 105 fath. (192 m.). (x 50.) 

Fig. 4. Phosphatised and ferruginous nodule, with large benthic Foraminifera. 
Stat. 5, Cape St. Blaize, 105 fath. (192 m.). (x 50.) 


PLatTE XXXV. 
Thin sections of nodules. 


Fig. 1. Glauconitic layer enclosing grains of glauconite (g) in the scoriaceous 
nodule of Pl. XXXITI, fig. 2. Stat. 7, Cape Point, 560 fath. (1024 m.). 
( x 60.) 

Fig. 2. Section of nodule formed of two rocks, showing line of contact. Stat. 10, 
Vasco da Gama, 166 fath. (304 m.). a, lower yellowish portion; 6, band 
strongly impregnated with glauconite and phosphate, in neighbourhood of 
contact; c, line of contact; q, quartz of upper portion. (x 45.) 

Fig. 3. Upper part of same nodule, showing the constitution of a phosphatised 
greensand nodule. gq, quartz; g, glauconite. (x 50.) 


XX XII. 


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‘DESCRIPTIONS o OF THE PALAEONTOLOGICAL MATERIAL 
“COLLECTED BY THE SOUTH AFRICAN Ss cn ee 


ee ee &. SEP 14 34 


5. me Contribution to ‘the Morphology of the Gorgonopsia. By 
ee _ Lieuwe D. Boonstra, D.Sc., Palaeontologist to the South 
= ee - - African Museum. oe Et Text- -figures. ) 


6. ‘Additions to our aencntletlie - the South African Soidinpsie: 
: preserved in the British Museum (Natural History). By 
a Lizuwe D. Boonstra, D.Sc., Palaeontologist to the South 
African Museum and Queen Victoria Scholar of the University 

| of cote (With 18 oe tenes) 
7 A Cais to the Morphology of the Mammal-like Reptiles 
et of the Suborder Therocephaha. By LinuwE D. Boonstra, 
BES. Palaeontologist ‘to the South African Museum and 
Queen Victoria Scholar of the iaaees of Stellenbosch. 
; (With 35 5 Text eee ) 


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5. A Contribution to the Morphology of the Gorgonopsia.—By Lizuwe D. 
BoonstrRA, D.Sc., Palaeontologist to the South African Museum. 


(With 11 Text-figures.) 


ALTHOUGH the first Gorgonopsian was described as long ago as 1860, 
and notwithstanding the fact that since then twenty-seven reputed 
genera and thirty-six species have been recorded from the South 
African Karroo Beds, our knowledge of these animals was until 
quite recently practically confined to their cranial morphology. 
The reason is not difficult to find. The Gorgonopsia were agile 
beasts of prey that lived on the higher and drier parts of the 
great Karroo Basin during Upper Permian and Lower Triassic 
times, so that on death their remains, in order to be entombed in 
the sediments, had to be transported to the lower-lying areas. 
During transportation the comparatively slender and more fragile 
bones of the postcranial skeleton were destroyed, whereas the more 
compact skull was more often preserved. Hitherto very few parts 
of the postcranial skeleton have been found in association with 
skulls. Owen, in 1876, described at length and gave good figures 
of the humerus of Cynodraco. Nothing more was added to our 
knowledge of the postcranial skeleton until Broom and Haughton, 
in 1913, jointly described and gave unsatisfactory photographs of 
the cervical vertebrae, pectoral girdle, and fore-limb of Aeluro- 
gnathus tigriceps. In a later paper, 1913, Broom gave a fuller 
description, with a figure, of the manus of Aelurognathus, and in 
the same year a short account of the shoulder-girdle of Scylacops. 
In 1921 Watson figured, without describing, the anterior cervicals 
of Scymnognathus whaitst. Then, in 1927, Pravoslavlev published 
two important papers on the Gorgonopsia of the North Russian 
Karroo Beds. Some of these forms had been photographed by 
Amalitzky, but had never been described. Unfortunately Pravo- 
slavlev’s work is in Russian, so that only the text-figures and photo- 
graphs are intelligible to most non-Russian scientific workers. It 
would appear, however, that in two genera—Inostrancemia and 
Amalitzkia—a practically complete vertebral column, pectoral and 
pelvic girdles, hind- and fore-limbs lacking only the feet, are known. 
In 1929 Haughton had the good fortune to obtain from the Rev. 
VOL. XXXI, PART 2. 15 


138 Annals of the South African Museum. 


J. H. Whaits a fairly well-preserved shoulder girdle and fore-foot 
of Lycaenodontoides. ‘These were fully described and figured. 

In a recent (1930) fairly comprehensive account of the Gor- 
gonopsia, Broom figured and described an imperfect vertebral column, 
pectoral and pelvic girdles, and the practically complete fore- and 
hind-limbs of Lycaenops. The known parts of Aelurognathus and 
Scylacops were also incorporated. 

During three collecting trips in the Southern Karroo I had the good 
fortune to collect a number of Gorgonopsians, which had considerable 
portions of the postcranial skeleton preserved. This material, 
augmented by the other previously described specimens preserved 
in the South African Museum, forms the basis of this communication. 
The collection consists of :— 

(a) S.A.M. Cat. No. 8950. Klein Koedoeskop, Beaufort West. 
Middle Tapinocephalus zone. 

A practically complete vertebral column, complete pectoral girdle, 
imperfect pelvic girdle, and fore- and hind-limb, lacking only some 
of the digits, are preserved. Unfortunately this specimen is preserved 
in a very intractable calcareous nodule, so that it has been very 
difficult to free the various skeletal elements from the matrix without 
more or less damaging the bone. The skull of this specimen has 
been described by Haughton under the name, Hipposaurus boonstrat. 
Collector, L. D. Boonstra. 

(6) S.A.M. Cat. No. 2342. Dunedin, Beaufort West. Low 
Cistecephalus zone. 

This specimen consists of a skull, seven cervical vertebrae, a com- 
plete though somewhat crushed pectoral girdle, a good humerus and 
radius, an imperfect ulna and a nearly complete fore-foot. This 
is the type specimen of Aelurognathus tigriceps described by Broom 
and Haughton. Collector, 8. H. Haughton. 

(c) S.A.M. Cat. No. 2348. Dunedin, Beaufort West. Low 
Cistecephalus zone. 

Associated with the skull there is an incomplete and crushed 
pectoral girdle, a good sternum and humerus, and some cervical 
vertebrae. This is the type of Scylacops capensis described by 
Broom. Collector, 8. H. Haughton. 

(d) S.A.M. Cat. No. 3329. Oudeberg, Graaff-Reinet. Cvste- 
cephalus zone. 

Associated with a snout, an imperfect pectoral girdle and manus 
‘ are preserved. Thisis Haughton’s type, Lycaenodontoides bathyrhinus. 
Collector, J. H. Whaits. 


A Contribution to the Morphology of the Gorgonopsia. 139 


(ec) S.A.M. Cat. No. 9344. Klein Bloemhof, Richmond. Ciste- 
cephalus zone. 

This specimen consists of a crushed skull; a continuous series of 
seven cervicals and seven dorsals (the rest of the posterior dorsals are 
lost), then a series commencing with the last presacral, three sacrals and 
five caudals; an excellently preserved pectoral girdle with a sternum; 
two good humeri, a good radius and ulna, and the proximal ends of 
the ulna and radius of the other side; an imperfect carpus; the left 
ilium and ischium and part of the right ischium; a good femur, tibia, 
fibula, tarsus, metatarsals, and some phalanges. For this specimen 
I propose the name—Aelurognathus microdon sp. nov. Collector, 
L. D. Boonstra. 

(f) S.A.M. Cat. No. 9345. Ou Plaas, Richmond. Cistecephalus 
zone. 

In this specimen there are preserved: a good, though slightly 
weathered and crushed, skull; a continuous series of seven cervicals 
and nine dorsals, together with some isolated dorsals and caudals; a 
nearly complete pectoral girdle with a sternum; two crushed humeri, 
a good right radius and ulna, and a crushed left radius and ulna; 
some isolated carpals, metacarpals, and phalanges. For this specimen 
I propose the name—Arctognathoides breviceps gen. et sp. nov. 
Collector, L. D. Boonstra. 

(g) S.A.M. Cat. No. 10188. Dunedin, Beaufort West. Low 
Cistecephalus zone. 

There is preserved a somewhat crushed skull with an excellent 
palate; a humerus, radius, and ulna; a tibia and fibula; and some 
crushed vertebrae. I have identified this as a specimen of Scylacops 
capensis, Broom. Collector, L. D. Boonstra. 


Skull. 


As the main object of this paper is primarily to note the increase 
of our knowledge of the postcranial osteology of the Gorgonopsia, 
no detailed account of the cranial morphology will be advanced. In 
the case of the three skulls of known species, only those points of 
structure which have hitherto not been fully understood will be 
mentioned. In the case of the two new forms the account will 


necessarily be somewhat fuller. 
Scylacops capensis (text-fig. 1, a, b, and c). 


The palate of the type (S.A.M. Cat. No. 3444) has been recon- 
structed by Broom (Mammal-like Reptiles, fig. 394). The anterior 


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Annals of the South African Museum. 


140 


A Contribution to the Morphology of the Gorgonopsia. 141 


part of the palate has now been partly exposed by removing part 
of the lower jaw, and it has now become clear that Broom’s recon- 
struction is at fault in a number of respects. The median vault 
formed by the pterygoid and palatine is as figured (fig. 1, c) and the 
prevomer tapers posteriorly, whereas Broom figured it as swelling 
out. Anteriorly the prevomer is wide with a median keel and a lateral 
flange. Unfortunately the palatal process of the premaxilla cannot 
be exposed without seriously damaging the whole snout. No inter- 
pterygoid suture is visible, but posteriorly there is a depression which 
may be an interpterygoid vacuity. As in Lycaenodontoides (fig. 2, a), 
there is a small foramen in the ectopterygoid. This bone extends 
right up to the antero-lateral corner of the transverse pterygoidal 
process. In my specimen (S.A.M. Cat. No. 10188), identified as 
Scylacops capensis, the palatine stretches very far forward and meets 
the palatal flange of the maxilla—a condition reminiscent of some 
Cynodonts (e.g. Glochinodontoides), Deinocephalians (e.g. Lamia- 
saurus), and Therocephalians (e.g. Whartsia). In my specimen the 
preparietal is a smaller and rounder bone than that of the type. On 
the ventro-posterior margin of the squamosal a thin wall-like flange 
of bone is developed, which appears to function as a partial external 
auditory meatus. In the zygomatic arch the relations of the 
squamosal and jugal are peculiar, in that the squamosal is clasped 
dorsally and ventrally by posteriorly directed forks of the jugal; 
laterally the squamosal appears as a ridge lying intercalated between 
the two jugal flanges. Neither in the type, nor in my specimen, 
does the occiput slope so much backwards as Broom figures it 
(Mammal-like Reptiles, fig. 34p). 


Lycaenodontoides bathyrhinus (text-fig. 2, a). 


Dr. Haughton has recently removed the matrix from the palate 
of his type-specimen (S.A.M. Cat. No. 3329), which now shows the 
structure of most of the anterior half. Two dentigerous ridges limit 
a fairly deep median vault, which is formed by both the pterygoid 
and palatines. There is no median pterygoid suture, but posteriorly 
a small vacuity is present. The ectopterygoids, which do not extend 
so very far down along the transverse pterygoid flanges, are pierced 
by a small foramen. The prevomers are separated from the ptery- 
goids by the palatines, which meet along a convoluted suture in the 
median line. In this it agrees with forms like Scylacops, Aelurosaurus, 
and Arctognathus. On the whole, the anterior part of the palate of 
Lycaenodontoides is very similar to that of Arctognathus curvimola. 


142 Annals of the South African Museum. 


Unfortunately it has not been possible to expose the extreme anterior 
end of the palate, so that nothing can be stated as to the nature of 
the palatal portion of the maxilla and premaxilla; but it would 
appear that the palate is on the whole a flat one, without the extra- 
ordinary vaulting shown by Arctognathoides (fig. 4, c). 


TEXT-FIG. 2. 


(a) Lycaenodontoides bathyrhinus. S.A.M. Cat. No. 3329. Ventral view of anterior 
part of the palate. x 4. 


(b) Arctognathus curvimola. Type. British Museum Natural History. Ventral 
view of anterior part of the palate. x about 3. 


For. =foramen in the ectopterygoid. 
Other lettering as in fig. 1. 


Arctognathus curvimola (text-fig. 2, b). 


Dr. 8. H. Haughton kindly let me copy a sketch of the palate of 
the type in the British Museum, and I quote the following from his 
notes: ‘‘My interpretation of the palate is entirely different from that 
of Watson. All the sutures are easily visible by wetting the skull; 
there is no interpterygoid suture; but there is a small interpterygoid 
vacuity. The arrangement of the bones is very similar to that in 
the form I have called Arctognathus whaitst. The sutures are all 
plainly visible as convolute lines—and different from the straight 
cracks which Watson has misinterpreted as sutures. The palate 
thus agrees with the Aelurosaurids in the separation of the pterygoids 


~ 


A Contribution to the Morphology of the Gorgonopsia. 148 


from the prevomers; but differs in the fact that the ectopterygoid 
does not extend on to the massive pterygoidal transverse process 
and in the nature of the ridge bounding the median vault.”’ There is 
thus much similarity between the anterior portions of the palates of 
Arctognathus and Lycaenodontoides, but there is some difference in the 
nature of the median vault and the proportions of the various elements. 

The dental formula in Arctognathus curvimola isi. 4, c. 1, m. 5, and 
in Lycaenodontoides 1.5, c.1, m.6; in the former, the molars are 
serrated. 


Aelurognathus microdon sp. nov. (text-fig. 3, a, b). 


The skull of this new species from Klein Bloemhof is somewhat 
crushed, and it has not been possible to expose the palate. Although 
it is much smaller than either A. tugriceps and A. serratidens it is 
undoubtedly cogeneric. The general configuration of the different 
bones is very similar, and it possesses a similar antorbital depression 
which is slightly overhung by the prefrontal; this depression is as 
deep as in A. serratidens, but deeper than in A. tigriceps. As in the 
other two species of Aelurognathus, the snout is high and rounded, 
and the width of the parietal region is much greater than the inter- 
orbital width, whereas, in the closely allied Scymnognathus, the 
width of the parietal region is relatively smaller. The temporal 
fossae are not wide in dorsal view, and the orbit does not look as 
much forward and outward as in the case of Scymnognathus. This 
species differs from the two hitherto known, mainly in that it is 
smaller; having in addition three and not four molars, which are also 
much smaller. The molars are not serrated, whereas in A. serratidens 
they are. Italso approaches A. tigriceps more closely in that the pineal 
foramen is small, whereas in A. serratidens it is large; it is situated 
further posteriorly than in both. The frontal forms a greater part 
of the upper orbital border than in A. tigriceps and a still greater part 
than in A. serratidens. The nasal stretches further posteriorly than 
in A. serratidens, but the frontal is larger than in A. togriceps. 


Arctognathoides breviceps gen. et sp. nov. (text-fig. 4, a, b, c, and d). 


Although slightly distorted by post-mortem crushing, it has been 
possible to prepare reliable dorsal, lateral, and ventral views of the 
skull. The most distinctive feature of this skull is its broad, short, 
and relatively high snout. In this it bears points of resemblance to 
Lycaenodontoides and Arctognathus. The interorbital region is, 
however, much wider than the parietal region, whereas in Arctognathus 


= = 


144 Annals of the South African Museum. 


it is actually slightly narrower. Furthermore, there does not appear 
to be such an overhang of the nasals over the nostrils. The nature 
of the palate is, moreover, totally different from that of these two 
genera. The massive transverse pterygoidal flanges are situated 
very far forward. Anterior to these flanges, the palatal roof con- 
tinues in anterior direction at a much higher level—there being an 


a 
SMe. Na, £¢ Pele Fr PF: POrb p, 


= SN 


TEXT-FIG. 3. 


Aelurognathus microdon sp. nov. S.A.M. Cat. No. 9344. x about 4. 
(a) Dorsal view of skull (crushing corrected). 
(6) Lateral view of skull a ws 


Ang. =angular. Art. =articular. Dent. =dentary. 
Other lettering as for previous figures. 


abrupt step of about 15mm. In the median line the anterior portions 
of the pterygoids and the postero-median portions of the palatines 
form a very curious rump, which, in its palatine portion, houses a 
median vault. Laterally and anteriorly there is yet another step 
of about 15 mm. on to the ventral surfaces of the ectopterygoids, 
prevomers, and antero-lateral portion of the palatines. Anterior 
to the transverse pterygoidal flanges, the palate thus contains two 
successive high vaults. The palate of Arctognathoides is thus very 


A Contribution to the Morphology of the Gorgonopsia. 145 


distinctive. In all other Gorgonopsians the palate is in comparison 
quite flat. The occiput does not slope backwards. The orbits look 
more outwards than upwards. The snout is square in section. The 


TEXT-FIG. 4. 
Arctognathoides breviceps gen. et sp. nov. S.A.M. Cat. No. 9345. x about }. 


(a) Lateral view of skull (crushing corrected). (c) Ventral view of skull (crushing corrected). 
(6) Dorsal view of skull 7) oe (d) Inner view of lower jaw. 
Cor. =coronoid. Qu.F. =quadrate foramen. Sur. Ang. =surangular. 


Pr.Art. =prearticular. Spl. =splenial. 
: Other lettering as in previous figures. 


infraorbital bar and zygoma are strong. The lateral ends of the 
paroccipital bone are directed posteriorly, so that the occiput is 
concave laterally. 

The preparietal is a small irregularly rounded bone. The pineal 
foramen is very small and situated far posteriorly. The frontals 
form about half of the upper orbital border. The orbit looks mostly 
outwards. The temporal fossa is large and roomy. 


146 Annals of the South African Museum. 


The dental formula is c.4,1.1,m.6. The molars are large; their 
posterior edges are distinctly serrated. There is a fairly distinct 
step in the dental border at the maxilla-premaxillary suture. The 
lower jaw is typically Gorgonopsian; the mentum is high, strong, and 
square; the coronoid process of the dentary is strong, but does not 
project into the temporal fossa as it does in Aloposaurus. The 
coronoid is large. There is one molar less than in the maxilla. 


The Chief Measurements. 


The measurements given here and in the subsequent tables are all 
taken by means of a Martin’s beam-compass, so that they all represent 
projections on to a vertical or a horizontal plane, given in millimetres. 


Aeluro- Arcto- Scylacops 
gnathus gnathoides capensis. 
microdon. breviceps. 


S.A.M. Cat. S.A.M. Cat. | S.A.M. Cat. 


No. 9344. No. 9345. No. 10188. 
Length : 
(Basioccipital-premaxilla) 197 187 156 
(Premaxilla-pineal fora- Ab 155 123 
men) 
(Pineal foramen-edge of 18 19 23 
occipital plate) 
(Snout—front of orbit) 105 90 84 
Length of lower jaw 1802 160 140 
Wrdth : 
(Across squamosals) 1402 150 120 
(Interorbital) 52 56 40 
(Intertemporal) 56 53 44 
(Across canines) 60 66 44 
(Across quadrates) 992 122 100 
Herght : 
(Maxillary edge—median . 63 (il 30 
suture) 
(Edge of occipital plate— 66 71 39 
basioccipital) 
(Quadrate—median 73 15 43 
suture) 
(Mentum of lower jaw) Q 52 33 
Length of molar series 9 35 i 


Length of diastema 22 12 10 


A Contribution to the Morphology of the Gorgonopsia. 147 


The Vertebral Column. 


Hitherto the complete presacral series was known in only one 
Gorgonopsian—lInostrancevia alexandert. This has been described by 
Pravoslavlev, but as his description is in Russian I am not able to 
appreciate the nature of his account, and, furthermore, am unable to 
count the number in the series from the photograph published. In 
Lycaenops, Broom found a continuous series of 20 vertebrae, and he 
believes that this comprises the complete dorsal series, and that, 
anteriorly, 7 cervicals were lost. Broom stated that this material is 
badly crushed, and he has been unable to determine the details of the 
various structural points. In Aelurognathus, Broom and Haughton 
and I have described the cervicals. 

In Hipposaurus boonstrar (S.A.M. Cat. No. 8950) I excavated the 
complete presacral, sacral, and probably also all the postsacral 
vertebrae. This is the first form from the Tapinocephalus zone where 
anything is known of the postcranial skeleton. This skeleton was 
preserved in a calcareous nodule which has proved to form an extremely 
intractable matrix, with the result that, during preparation, the 
caudals have been seriously damaged and their sequence lost, and 
even in the presacrals many of the processes have been damaged. 
In the presacral series there are 28 vertebrae. Of these 7 are cervicals, 
15 typical dorsals, and the posterior 6 are perhaps best referred to as 
lumbar vertebrae. There are 3 sacrals and at least 26 caudals. The 
total length of the presacral series is 700 mm.; of the sacrals 55 mm., 
and of the caudals at least 450 mm. 

In Aelurognathus microdon (S.A.M. Cat. No. 9344, fig. 5, a, b) there 
is present an excellently preserved presacral series of 14 vertebrae, of 
which the anterior 7 are cervicals; then, probably 12-13 vertebrae 
are missing, and then there is a further continuous set commencing 
with the last dorsal, which includes 3 sacrals and 5 caudals. 

In Arctognathoides breviceps (S.A.M. Cat. No. 9345, fig. 5, c, d) a 
continuous series of the first 16 presacrals and a further number of 
disarticulated dorsals and caudals are preserved. These vertebrae 
are all slightly crushed, and in most the tops of the neural spines have 
suffered from weathering. 


The Cervicals. 


I have already described the cervicals of Aelurognathus tigriceps 
(fig. 5, e) in a short note—“ The Cervical Vertebrae of a Gorgonopsian 
(Aelurognathus tigriceps)’’ in the Ann. Mag. Nat. Hist., Series 10, vol. 


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A Contribution to the Morphology of the Gorgonopsia. 149 


xil., 1934. The excellent material now at my disposal fully confirms 
the views expressed in this paper. In Hipposaurus, Aelurognathus, 
and Arctognathordes the neck consists of 7 vertebrae. In Hipposaurus 
and Aelurognathus the _ shoulder- 
girdle is preserved in natural relation 
to the vertebral column so that this 
important point in the determina- 
tion of the cervical series is known. 
In addition the presence of inter- 
centra in the first 7 vertebrae, the 
abrupt increase in length of the 
diapophyses in the 8th vertebra, the 
upward shifting of the parapo- 
physial facet, and the change of the 
zygapophysial articulation from a 
horizontal plane to one at 45°, are all 
points that definitely show that the 
neck consisted of 7 segments. In 
all three genera a pro-atlas is pre- 
served, and has the relations as 
described for A. tugriceps; the atlan- 
tal arches composed of two unfused 
halves are essentially the same in 
the three genera, although they 
differ somewhat in point of shape 
and size; the large atlantal inter- 


Pr Atv 


centrumand untfused atlantal pleuro- 
centrum appears to be constant in 
the Gorgonopsia. In all three 
genera the atlantal dichocephalous 
rib articulates with a diapophysis on 
the atlantal arch and with a parapo- WO se arcns 
physial facet on the atlantal inter- S.A.M. Cat. No. 2342. 


centrum. In the succeeding verte- Dorsal view of the first 6 vertebrae. 
x about $. 


4 


{bapa 5), @ 


brae the parapophysial facet shifts 
on to the extreme anterior edge of epee ee 
the centrum, and the diapophysial facets are carried on strong 
ventro-posteriorly directed processes. The zygapophysial articula- 
tions are horizontal. All the centra are deeply amphicoelous— 
practically notochordal. The neural spine of the axis is typically 


differentiated as in all Therapsids. The shape differs somewhat in 


150 Annals of the South African Museum. 


the various genera (fig. 5,a,c,e). Only in Hipposaurus do the cervicals 
carry a strong ventral keel to the centra. In A. tigriceps the spine of 
the 3rd vertebra appears to be rudimentary. This may be patho- 
logical, as in none of the others does this condition obtain. 
Professor D. M. 8. Watson has in his collection at University College 
a skull and a series of 8 cervical vertebrae of a specimen of Scylacops 
capensis from Wellwood, Graaff-Reinet, which had been presented to 
him by the South African Museum. The cervical series is in a fairly 
good state of preservation, and confirms the conclusions arrived at in 


Text-ria. 5, f. 
Aelurognathus microdon. Ribs. x about 4. 
1. Lateral view of the axial rib. 


2. ee = 4th rib (proximal end). 
oe 9 ” 7th 33 9 ” 
4. be) 9° 12th 99 99 99 


our study of the neck in Aelurognathus, Hipposaurus, and Arcto- 
gnathoides, viz. the structure of the pro-atlas, atlantal arches, the 
odontoid, and the atlantal intercentum is as described in those forms; 
the same evidence of the presence of 7 cervicals is present. The 
anterior cervicals of Scylacops agree with those of Hipposaurus in that 
they also have a slight ventral keel developed on the centra. 
Watson’s skull shows a very good articulation between the basi- 
sphenoid and the posterior ramus of the pterygoid; the stapes and the 
foramen for the 10th nerve are beautifully preserved. The frontal 
seems to take a greater part in the formation of the orbital border 
than is shown in the two specimens in the South African Museum. 


The Dorsals. 


In Hipposaurus, where the dorsal series is fully preserved, there are 
21 vertebra; in Lycaenops, Broom maintained that the 20 elements 


A Contribution to the Morphology of the Gorgonopsia. LID 


preserved comprises the whole dorsal series. In Hipposaurus the first 
15 elements of the dorsal series are typical dorsal vertebrae. Con- 
tinuing backwards from the 22nd presacral to the first sacral, the 
vertebrae decrease considerably in height and width, becoming thereby 
relatively longer and more slender; the diapophyses also become more 
slender and weaker, and no indication of a facet for the capitulum of 
the ribs can be seen, and thus the possibility arises that the posterior 
ribs were either single-headed or that these vertebrae are really lumbar 
and possessed no ribs; it is, moreover, remarkable that the first 15 
dorsals possess no intercentra, whereas in the posterior presacrals 
intercentra are developed, nearly as strongly asinthe neck. Although 
Broom, in his reconstruction of Lycaenops (Mammal-like Reptiles, 
fig. 45), figured ribs right up to the sacrum, I am inclined to think that 
in the Gorgonopsia the vertebrae immediately anterior to the sacrum 
carried noribs. This would be in harmony with the rest of the skeleton, 
which indicates fairly lightly built animals of considerable agility. The 
absence of ribs in the lumbar region would greatly increase the flexi- 
bility of the back—particularly in lateral direction. If I am correct 
in this point of view the Gorgonopsia would in this respect approach 
the condition of some Cynodonts (e.g. Thrinaxodon), Cotylosaurs (e.g. 
Inmnoscelis, Labidosaurus), and Pelycosaurs (e.g. Sphenacodon). 

In the three genera—Hipposaurus, Aelurognathus, and Arctogna- 
thordes—the width across the zygapophyses in the dorsal vertebrae 
abruptly decreases at the transition from the cervicals; the diapo- 
physial processes increase in length and are more horizontally 
directed, so that the facet is at a higher level than in the cervicals; 
the parapophysial facets shift higher up on to the centrum, and in 
Hipposaurus are not seen after the 15th dorsal. Continuing back- 
wards the centra progressively increase in length. Whereas in the 
cervicals the zygapophyses are horizontal, in the dorsals they lie at 
45° to the horizontal, which indicates that less lateral movement was 
possible. The centra are deeply amphicoelous, with a moderate 
lateral constriction; ventrally the centra are concave in antero- 
posterior direction, but are not pleurocoelous; no keel is developed. 
In Aelurognathus microdon, where the neural spines are perfectly 
preserved, they are very tall and laterally compressed, and are very 
similar to those of Amalhtzkia annae figured by Pravoslavlev; in 
A. tagriceps the cervical spines are short and much thicker; in Arc- 
tognathoides the spines are shorter than in A. microdon and slightly 
less laterally compressed; in Hipposaurus they were apparently 
shorter still, and also somewhat laterally compressed. 


152 Annals of the South African Museum. 


Sacrum. 


The sacrum is well-preserved in A. microdon, but is somewhat 
crushed in Hipposaurus. In the former, three vertebrae contribute 
to the support of the ilium. Their centra are fairly firmly, although 
not completely, fused. Each carries a strong sacral rib which emerges 
from rather high up on the middle of the centrum. Distally the ribs 
spread out fan-wise to abut deeply on the ilium. The first rib is the 
strongest, and its distal end stretches very far ventrally—to below the 
upper half of the acetabulum. The posterior two abut higher up on 
the ihum. As in Mammals, the ribs abut on the postero-internal 
surface of the ilium, leaving the anterior part unsupported. In 
Hipposaurus the upper portion of the crest of the left ilium is pre- 
served in its natural position. Here also 3 sacral ribs support it— 
abutting on the postero-internal surface. Broom’s inferences, as to 
the condition in Lycaenops, appear, in the light of this new material, 
to be substantially correct. 


Caudals. 


In no specimen are the caudals fully preserved. In Hipposaurus 
there are at least 26. In Aelurognathus microdon the first 5 caudals 
are preserved. The neural spines are lower, shorter, and less com- 
pressed than in the dorsals, and the transverse processes are strong 
and extend as far laterally as the external iliac surface. Judging by 
the imperfectly preserved tail of Hipposaurus, it would appear that in 
the Gorgonopsia the tail was long and whip-lke. 

The chief measurements are: 


Vertebral Column. 


Aeluro- Hippo- Aeluro- Arcto- 
gnathus SQUTUS gnathus | gnathoides 
tigriceps. | boonstrai. | microdon. | breviceps. 


S.A.M. S.A.M. S.A.M. S.A.M. 
Cat. No. | Cat. No. | Cat. No. | Cat. No. 


2342. 8950. 9344. 9345. 
Length of 7 cervicals. 270 191 167 180 
Ist 7 dorsals : 2 155 176 200 
Height of Axis . 76 37 57 43 
_ 3rd cervical : 56 32 67 4] 
= 4th ,, : a 35 71 47 


a Ist dorsal . : Q om 81 5T 


A Contribution to the Morphology of the Gorgonopsia. 153 


Vertebral Column—continued. 


Aeluro- Hippo- Aeluro- Arcto- 
gnathus SQUTUS gnathus | gnathoides 
tagriceps. | boonstrat. | microdon. | breviceps. 


S.A.M. S.A.M. S.A.M. S.A.M. 
Cat. No. | Cat. No. | Cat. No. | Cat. No. 


2342. 8950. 9344. 9345. 
Width across Diapophyses : 
(a) Axis : : 60 34 34 41 
(b) 3rd cervical. 61 36 36 45 
(yeoth %, 74 38 37 47% 
@)xith ~~; 84 40 44 48 2 
(e) 1st dorsal : : Q 43 46 49 
(i) Sle : ! 2 45 50 51 
Length of Centra : 
(a) Axis : 32 26 21 21 
(b) 3rd cervical . 33 27 23 22 
@ycth? 4 ‘ : Q 22 24 20 
(d) 5th dorsal q 22 27 21 
Width across Prezygapophyses: 
(a) Axis 34 25 20 29 
(b) 3rd cervical. ; 35 30 22 27 
(GG) ed ee : 4 32 24 22 28 
(d) 5th dorsal : q a 21 23 
Width of Centrum : 
(a) Axis ‘ 27 16 20 26 
(b) 7th cervical. 34 15 21 23 
(c) 5th dorsal a 14 20 17 


The Ribs (fig. 5, f). 


The ribs in all the specimens are very badly preserved. The 
proximal ends, however, are preserved in quite a number of cases. 
In all, the dorsal ribs are long and slender—those of Hipposaurus are 
not more than 4 mm. thick; in Aelurognathus, however, they are some- 
what thicker (12 mm.). In A. mcrodon some of the cervical ribs are 
preserved—the axial rib is a short, wide, and leaf-like bone, the 4th 
is already long and hasa slender shaft. All the ribs that are preserved 
are dichocephalous, the tuberculum and capitulum being separated 

VG sexx PART 2 16 


154 Annals of the South African Museum. 


by a slight notch; it is, however, possible that the posterior dorsals 
are single-headed. In no case has it been possible to determine the 
nature of the junction of the anterior dorsal ribs with the sternum. 
Broom, in his reconstruction (1930) of the pectoral girdle of Aeluro- 
gnathus tigriceps, indicated three ribs articulating with the sternum. 
There is, however, no evidence that this is so, although in the sternum 
of Lycaenops he maintained to have noticed three notches on the lateral 
edge of the sternum, to which an equal number of ribs apparently 
articulated. 


The Pectoral Girdle (text-fig. 6). 


The Gorgonopsian breast-shoulder-apparatus is now very well 
known. It is fully preserved in Hipposaurus (from the Tapino- 
cephalus zone), in Aelurognathus microdon, in Arctognathoides 
breviceps, and somewhat incompletely and crushed in Aelurognathus 
tigriceps, Lycaenodontoides and Lycaenops. In the Russian forms 
Pravoslavlev has described it in Inostrancevia and in Amalitzkia. It 
consists of a scapular part formed by the three cartilage bones— 
scapula, coracoid, and precoracoid—and a clavicular part, which con- 
sists of three dermal bones—clavicles, cleithra, and an unpaired inter- 
clavicle; in addition an ossified sternum is always developed. The 
scapula is a tall and slender bone with the dorsal part of its blade con- 
siderably expanded; it carries no acromion process, and thus no supra- 
spinatus fossa as in mammals and closely related reptiles; it forms half 
of the simple, posteriorly directed non-“ screw-shaped”’ glenoid cavity ; 
on its ventro-anterior end an extensive plate of bone is developed, 
which received the m. supracoracoideus. There is no supraglenoid 
foramen, but internally the scapula forms the dorsal border of the 
strongly developed subscapular fossa. In the different genera the 
scapula is very similar, except that in the Russian forms the dorsal 
expansion of the blade appears to be relatively greater than in the 
South African forms. 

The greater part of the coracoid plate is formed by the procoracoid, 
which is tall and long (particularly in Hipposaurus); the procoracoid 
foramen is situated immediately anterior to the glenoid and passes 
through the bone in dorsal direction; the procoracoid just enters the 
glenoid cavity to form a small part of its extreme anterior border; 
the coracoid has a well-developed posteriorly developed lip, from 
which the coraco-brachialis and costo-coracoideus arose; anteriorly 
the coracoidal plate is closely applied to the internal face of the 
clavicle, and ventrally it rests on the median stem of the interclavicle. 


Cor. 


TExtT-FIG. 6.—Pectoral girdles. 


(a) Hipposaurus boonstrai. S.A.M. Cat. No. 8950. Ventral view of interclavicle. 
x 2. 
(6) Hipposaurus boonstrai. S.A.M. Cat. No. 8950. Lateral view. x 2. 


(c) Aelurognathus microdon sp. nov. S.A.M. Cat. No. 9344. Ventral view of 
sternum. xX #3. 


(d) Aelurognathus microdon sp. nov. S.A.M. Cat. No. 9344. Lateral view. x i. 
(e) = be - a Internal view. x i, 
Cl. =cleithrum. P.Cor. = procoracoid. 
Cla. =clavicle. P.Cor.For. = procoracoidal foramen. 
Cor. =coracoid. Se. = scapula. 


I.Cl. =interclavicle. St. = sternum. 


S Nees oF 
S ea ae 5 Z Zz LZ 
SSEZ2Z 
StS S22 


TExtT-FIc. 6.—Pectoral girdles—continued. 


(f) Arctognathoides breviceps gen. et sp. nov. S.A.M. Cat. No. 9345. Lateral view. 3. 
(9) 23 39 44 a Internal view. x#. 
(h) + 99 5p a) Posterior view. x. 
(1) Aelurognathus tigriceps. S.A.M. Cat. No. 2342. Lateral view. x 4. 

(4) Scylacops capensis. S.A.M. Cat. No. 2343. Lateral view. x }. 

(k) 99 oS 50 55 Ventral view of sternum. x }. 

(Z) > » 9 55 Ventral view of left humerus. x#. 


Lettering as on previous page. 


A Contribution to the Morphology of the Gorgonopsia. 157 


In the older Hipposaurus the plate appears relatively larger than in 
the forms from the Endothiodon and Cistecephalus zones. 

The clavicle is a large bone, being particularly massive in Lycae- 
nodontoides; dorsally it has a long process which is applied, very 
high up, to the antero-internal edge of the scapula; ventrally it has 
a triangular expansion with a relatively long base; this clasps the 
lateral expansions of the anterior end of the interclavicle. In A. 
tigriceps, A. microdon, and Lycaenodontoides the ventral ends of the 
clavicles nearly meet and thus hide the anterior end of the inter- 
clavicle from ventral view; in Hipposaurus, however, the clavicles 
are widely separated by the interclavicle (fig. 6, a). 

The interclavicle is a large bone, as long as the coracoidal plate; 
anteriorly it is expanded, and lies nearly wholly internal to the 
expanded ends of the clavicles, except in Hipposaurus, where in 
ventral view it is shovel-shaped; posteriorly it is shaft-like for a 
short distance and then expands to form a long oval extremity, 
which, lying ventrally to the sternum, supports that element along its 
anterior half. In Lycaenodontoides, Arctognathoides, and Aelurognathus 
microdon a strong keel, which is developed on its medio-ventral 
surface, received the pectoral muscle; in A. tegriceps and Hippo- 
saurus, however, there is only a slight ridge on the medio-ventral 
surface. 

The sternum is present in the type-specimens of Scylacops capensis, 
Lycaenops ornatus, Aelurognathus microdon, Arctognathoides, and, 
in part, in Hipposaurus. In Scylacops (fig. 6, k) it is roughly square, 
whereas in all the others it is roughly oval in outline. In Lycaenops, 
Broom figured (Pl. XX VIII, 18, 1930) the sternum as an elongated bone 
which has three notches on either side for the reception of the first 
three dorsal ribs. In the sterna examined by me no such definite 
notches have been noticed, but it is possible that the sternal edges 
were cartilaginous, and the junction with the ribs would in that case 
not be preserved. 

The chief measurements are: 


[TABLE 


158 Annals of the South African Museum. 
Pectoral Girdle. 


Hippo- Aeluro- Arcto- | Scylacops | Aeluro- 
SQUTUS gnathus |gnathoides| capensis. | gnathus 
boonstrai. | microdon. | breviceps. tigriceps. 


S.A.M. S.A.M. S.A.M. S.A.M. S.A.M. 
Cat. No. | Cat. No. | Cat. No. | Cat. No. | Cat. No. 


8950. 9344. 9345. 2343. 2342. 
Height : 
Total coraco- 113 178 155 128 230 ? 
scapular 
Glenoid edge-top 99 123 116 98 130? 
of scapula 
Greatest procora- 422 57 43 36 2 115 
coidal 
Greatest clavi- 78 107 1002 Q 163 
cular 
Length : 
Coracoid plate 77 90 75 55 2 143 
Interclavicle 105 103 81? 2 210 
Sternum 25 54 61 +4 Q 
Blade of scapula 24 52 4] 33 65 2 


Fore-limb (figs. 7-9). 

As a whole the fore-limb is fairly light and slender. In Hippo- 
saurus the relative length and slenderness of the limb is much greater 
than in Cynodraco, Aelurognathus, Arctognathoides, Scylacops, Ino- 
strancevia, and Amalitzkia. But, notwithstanding the relative 
slenderness, the humerus still retains most of the typically primitive 
reptilian characters, viz. (although constricted in its middle, one can 
hardly speak of a humeral shaft as it is known in mammals) both 
the distal and proximal ends are expanded and are twisted around 
the “shaft”? to subtend an angle of about 40°; the proximal articu- 
latory surface is elongated, and, although mainly articulating with 
the glenoid cavity with its pre-axial corner, there is not much 
approach to the typical ball-and-socket joint of mammals; the glenoid 
faces backwards and downwards, so that the humerus no longer 
stands out practically horizontally as in the primitive Cotylosaurs; 
the animal, however, does not stand vertically on the fore-limbs, but 
still hangs somewhat between its supports; the distal articulation 


A Contribution to the Morphology of the Gorgonopsia. 


159 


for the radius and ulna is much more terminal than in Cotylosaurs 
and Pelycosaurs, and the trochlear fossa is quite deep, so as to allow 


considerable extension of the epipodial; 
the gait of the Gorgonopsia was thus 
much more upright than in the more 
primitive reptiles; the delto-pectoral 
crest 1s fairly strong and extends far 
distally; it is, however, much weaker 
than in Cotylosaurs, Pelycosaurs, Deino- 
cephalians, and the higher Cynodonts 
(Cynognathus); in particular it appears 
that the pectoral muscle was much 
weaker than in the forms mentioned; 
the delto-pectoral flange is demarcated 
from the dorso-proximal surface by a 
weakanteriordorso-ventral line; distally, 
the confluent ectepicondylar and sup- 
inator flange is strong, indicating strong 
extensor muscles; on the post-axial side 
the entepicondylar flange is also quite 
well developed, indicating strong flexors. 
Except in point of size there appears to 
be very little difference in the humeri 
of the known forms of the Gorgonopsia 
—Hipposaurus is distinct in possessing 
a long and slender humerus; in Arcto- 
gnathoides the epicondyles are more 
strongly developed than in the others; 
the keel on the interclavicle in Lycae- 
nodontoides, Aelurognathus, and Arcto- 
gnathordes indicates a greater degree of 
localisation of the pectoralis than in 
the geologically older Hipposaurus. 
Proximally, the radius fits fairly firmly 
in the sigmoid notch of the ulna; proxi- 
mally, the chief articulation of the 
eplpodial is formed by the radius, which 
fits on to the rounded capitellum of the 
humerus; the ulna has a trochlear joint, 


H 
TExtT-FIG. 7. — Hipposaurus 
boonstrat. S.A.M. Cat. No. 
8950. Dorsal view of part of 
the left fore-limb. x 4. 


c2 =first and second central. 
= humerus. 
=intermedium. 

=radius. 

=radiale. 

=ulna. 

=ulnare. 

= distal carpals. 

= metacarpals. 


le) 
-— 
“ 


fh ri 


LL 
ao 


and its olecranon fits into a 


deep trochlear fossa when the lower limb is extended; distally, the 
intercalation of the intermedium further prevents much. independent 


TEXT-FIG. 8. 


Aelurognathus microdon sp. nov. S.A.M. Cat. No. 9344. 
(a) Dorsal view of right humerus. x }. 
(6) Ventral view of right humerus. x }. 
(c) Anterior view of left fore-limb. x i. 


Arctognathoides breviceps gen. et sp. nov. S.A.M. Cat. No. 9345. 


(dz) Dorsal view of right radius and ulna. x #4. 
(e) Ventral view of right radius and ulna. x 4. 


Scylacops capensis. S.A.M. Cat. No. 10188. 
(f) Dorsal view of right tibia and fibula. x 4. 


(g) Anterior view of right humerus, radius, and ulna. x 
el, c2 =first and second central. R. =radius. 
D.P.C.R. =delto-pectoral crest. r. =radiale. 
ect. =ectepicondylar foramen. U. —uina: 
ent. =entepicondylar foramen. u. =ulnare. 

Fie =humerus. 1-5 = distal carpals. 


ac =intermedium. 


ib 
ze 


A Contribution to the Morphology of the Gorgonopsia. 161 


movement by either radius or ulna. The proximal post-axial border 
of the ulna is rugose for a firm insertion of a strong triceps muscle. 
The Gorgonopsian manus is known from a number of forms— 


DPCR. 


TEXT-FIG. 8—continued. 


Cynodraco major. B.M.N.H., 47310. 
(hk) Ventral view of left humerus. x 4. 


Lettering as on previous page. 


Aelurognathus tigriceps, A. microdon, Lycaenodontoides bathyrhinus, 
Lycaenops ornatus, and Hipposaurus boonstrai—in which it is either 
partly, or, in two cases, practically completely preserved. The 
carpus consists of three rows of elements and has the formula 3, 2, 
4 or 5; in the proximal row there are radiale, intermedium, and 


162 Annals of the South African Museum. 


ulnare, with a space between them for the perforating carpal canal; 
in the central row two well-developed centrals are present; in the 


/ fs 7 

Hh 
I, bef 
“4 pi 

RCH) 


I; 
[4 


I / 
/ Hh | 
WY [& 
d KT 
/fi 
/ WY, 
Le AL 
an 4 / 
! WW, [4 
\ ia 


5) 


TEXT-FIG. 8—continued. 


Cynodraco major. B.M.N.H., 47310. 
(2) Dorsal view of left humerus. x 3. 
(j) Preaxial view of left humerus. x 4. 


Lettering as on page 160. 


distal row there are 4 bones in Aelurognathus, Lycaenops, and Hippo- 
saurus, but in Lycaenodontoides the 4th and 5th distals are not fused, 
so that here 5 bones are present; this was noted by Haughton in 
Lycaenodontoides, and I have re-examined the type-material and 
find that there is definitely an unfused 5th distal; in Lycaenops, 


TEXtT-FIG. 9.—Aelurognathus tigriceps. S.A.M. Cat. No. 2342. x i. 


(a) Anterior view of left fore-limb. (6) Ventral view of left humerus. 
cl,c2 =first and second central. p- =rudiment of a prepollex (radial 
D.P.C.R. =delto-pectoral crest. sesamoid). 
ect. =ectepicondylar foramen. r. =radiale. 
ent. =entepicondylar foramen. u. =ulnare. 

i. =intermedium. 1-5 =distal carpals. 


I-V = metacarpals. 


164 Annals of the South African Museum. 


Broom has recorded a fully preserved pisiforme, and in Aelurognathus 
tigriceps a piece of bone representing the pisiforme is preserved. In 
no Gorgonopsian is the whole acropodium preserved; in A. tigriceps 
4 digits are preserved, in Lycaenops 4 digits, some lacking the claw 
phalanx, and in Lycaenodontoides only the 3rd and 4th digits are 
complete. In A. tigriceps, Broom gave the formula 2, 3, 4, 5, 3, and 
subsequently the same formula was advanced for Lycaenops; 
Haughton has given the formula ?, 2, 3, 3, ? for Lycaenodontoides, 
and an examination of this material shows that Haughton’s figures 
are correct. It is only recently that I have adequately removed the 
matrix from the beautifully preserved manus of Aelurognathus 
tigriceps. ‘The metacarpals are as figured (fig. 9, a); they have 
curiously formed ends; distally, in particular, they look very much 
like epiphyses; the 5th is of complex shape, the 4th squat, and the 
others elongated; the phalangeal formula I find to be 22, 3, 4, 4, 3. 
In the 4th digit the second phalanx is deeply constricted, but very 
careful preparation has revealed that it is a single bone, and not two 
separate elements as Broom thought. The question now arises 
whether Broom is correct in figuring two short elements as the 2nd 
and 3rd phalanges of the 4th digit in Lycaenops. I have not seen 
this material, but on the facts at my disposal I think that in Lycaenops 
the 4th digit only possessed 4 segments. Both in Lycaenops and 
A. tigriceps the 2nd phalanx of the 3rd digit is very short, and it 
appears probable that in this digit the number of phalanges is under- 
going reduction. In this case it is of great interest to note that in 
Lycaenodontoides both the 3rd and 4th digits possess only 3 segments. 
The chief measurements are : 


2 = ea 
S 8 ol8 chalS cdisle cdelS edaleess 
Ԥ 3 3 Qi. So Q.2 cS 0 
SSS |Se sls scSls coh Seeals ccs 
Sse (esa (88a sa. (See ssa 
SS48/8 84 5/SFs S/S S48 |S S< 8 |F S45 
SS es ee | Sis aa 
NX 
Length of : 
Humerus : : : : 174 159 144? aa 226 90? 
Radius . : ; ‘ : 12% lll 101 ? 162 83 
Ulnay = . : : 3 147 123 Bel ? 200? 93 
Width across : 
Epicondyles of humerus . : 44 57 75? 50 113 35 
Proximal expansion of humerus 47 61 60 43 121 33? 
Minimum of humeral shaft A 16 17 al 12 32 14 
Proximal end of radius . : 23 28 3) ? 51 18 
Distal end of radius ; 3 ile 24 25 ? 44 17 
Proximal end of ulna a a 31 38 37 ? 65 24 
Distal end of ulna . ? ? 


A Contribution to the Morphology of the Gorgonopsia. 165 


TExt-FIc. 10.—Aelurognathus microdon sp. nov. S.A.M. Cat. No. 9344. 


(a) Lateral view of left half of the pelvis. x 4. 
(6) Left femur, ventral. x 4. 
(@) seers ba dorsal. x 4. 

»» posterior. x 4. 


(d)_,, 
(e) Left hind limb, dorsal. x 4. 


F. =femur. t. =tibiale. 

f. =fibulare. Tr. Ex. =external trochanter. 
Fi. =fibula. Tr.In. =internal trochanter. 
1. =intermedium., 1-5 = distal tarsals. 


a vilbiar IV =metatarsals. 


166 Annals of the South African Museum. 


Pelvic Girdle (fig. 10, a). 

Hitherto the pelvis of the Gorgonopsia was known in only two 
forms—Lycaenops ornatus and Inostrancevia alexandert. In the 
former only the ilium is perfectly preserved—both the pubis and 
ischium have parts missing. In Aelurognathus microdon the left ilium 
is perfect, whereas on the right side only the internal face with the 
distal ends of the three sacral ribs in position is well shown; the left 
and part of the right ischia are preserved; only the acetabular portion 
of the right pubis is preserved. The ilium is vertical, long, and low, 
and forms more than half of the acetabulum; the main sacral sus- 
pension is pre-acetabular; the anterior process of the iliac blade 
extends far anterior to the acetabulum (strong ilio-femoralis) whereas 
the posterior process is in line with the posterior edge of the aceta- 
bulum (tail extensors of medium strength); the acetabulum is directed 
outwards. The ischium is long, but the ischial tuberosity does not 
appear to be strong; the symphysis is weak, but the upper edge is 
rounded and thickened; it forms less than + of the acetabulum, and is 
not strongly fused with the illum. The pubis forms considerably less 
than 1 of the acetabulum; its articulation with the ium and ischium 
is weak; itis of great antero-posterior length—ain this respect differing 
greatly from Deinocephalians and Cynodonts. In Hipposaurus the 
crest of the right ilium is preserved, and on the left the ischium and 
pubis and acetabular portion of the ilium are preserved. Unfortun- 
ately, however, the pes overlies most of the left ischium and cannot 
be removed without destroying it. The ischium and ilium do not 
differ greatly from that of Aelurognathus. The ilium of both these 
genera differ from that of Lycaenops, where the anterior and posterior 
iliac processes are of equal length. In the pubis of Hipposaurus the 
foramen is not encircled by bone, but 1s really a notch in the pubis to 
which the ischium forms the posterior border; medially it appears to 
be open, and appears to be confluent with its fellow of the other side; 
unfortunately the specimen is not very well preserved, so that some 
uncertainty exists. In Lycaenops the foramen is also not encircled by 
bone in the specimen as preserved, but Broom thought that in life a 
typical pubic foramen was present. In view of the known condition 
in Hipposaurus, it would appear that the Gorgonopsia possessed a pubo- 
ischiadic fenestra and no pubic foramen (homologous to the “ obturator 
foramen”? of Mammals, which includes the real obturator foramen 
merged with the pubic foramen). The upper border of the pubis is 
very much thickened in Hipposaurus, and from this shaft-like edge a 


A Contribution to the Morphology of the Gorgonopsia. 167 


thin flange of bone proceeds medially to meet its fellow in a very weak 
symphysis. 
The chief measurements are: 


Hipposaurus | Aelurognathus 
boonstrar. microdon. 


S.A.M. Cat. S.A.M. Cat. 


No. 8950. No. 9344. 
Length ofilium .. : 64 102 
ischium . , é ; 65 82 
ee) pubis fk ; . : 56 Q 
ue acetabulum . : : 32 A] 
Height of ilium from the acetabular 
border : ! ¢ 49 


Hind Limb (figs. 10, 11). 


In the American Museum of Natural History the pelvis and both 
hind limbs of a specimen referred by Broom to ? Aelurosaurus felinus 
is preserved. The femur has been figured by Williston (Osteology of 
the Reptiles, fig. 132, 6). In Lycaenops the hind limb has been 
shortly described and only an indifferent figure of the limb has been 
published, together with a reconstruction of the pes. In Aelurogna- 
thus microdon a beautiful hind limb, lacking only the distal phalanges, 
is preserved. In Hipposaurus boonstrai the complete right hind limb 
was preserved, but has unfortunately been damaged in preparation 
due to the intractable nature of the matrix. 

The hind limb is appreciably longer than the fore limb, and as its 
elements, particularly the femur, are much more slender, an appear- 
ance of still greater lengthis created. The femur isa long slender bone 
with a long shaft and unexpanded ends, which are only slightly 
“twisted”? on the shaft; the proximal end is directed preaxially, and 
its articulatory face is rounded (it is more elongated in ? Aelurosaurus) ; 
There is no “neck,” but there isa slight notch on the proximal post- 
axial edge between the proximal surface and the external (major) 
trochanter; this trochanter is situated far proximally, and continues 
distally as a narrow flange for the gluteal muscles; on the ventro- 
postaxial surface a low ridge represents the internal trochanter, which 
is much less prominent than in Therocephalians and Cynodonts; there 
is no intertrochanteric fossa, and the m. pubo-ischio-femoralis externus 


168 Annals of the South African Museum. 


must have been weak; distally the condyles are not separated by a 
deep intercondylar sulcus; on the ventro-distal surface there is a 
fairly deep popliteal fossa; on the dorso-distal surface there is a slight 


Text-Fic. 11.—Hipposaurus boonstrai. S.A.M. Cat. No. 8950. Right 
hind limb, dorsal view. x 4. 


Lettering as in previous figure. 


depression, so that it would appear that the femora-tibial muscle was 
weak. It is evident from the points raised in the structure of the 
femur that in the Gorgonopsia the body was not slung in between the 
hind limbs, but, on the contrary, the gait must have been more or less 
upright, and the locomotory movements executed with considerable 


A Contribution to the Morphology of the Gorgonopsia. 169 


agility. The epipodial, though long and slender, is shorter than the 
propodial; the fibula has a long thin shaft with relatively greatly 
expanded ends, and it has a strong postaxial curvature, and as the 
tibia also had a slight preaxial curvature the spatium interosseum is 
great; the tibia has a widely dorso-ventrally expanded proximal end, 
but its distal expansion is weak; its cnemial crest is not strongly 
developed; proximally both elements articulate with the distal end 
of the femur; the articulation of the tibia is the stronger, but the 
fibula directly receives the femur and does not form a sliding joint on 
to the postaxial femoral epicondyle; distally the articulation with the 
tarsus is not a very firm one—a considerable amount of cartilage must 
have been developed. The tarsus of Lycaenops has been described 
by Broom; he found that the epipodial articulated with only two 
elements in the proximal row of the tarsus; these he thought were the 
astragulus and caleaneum; distally, on the radial side a navicular 
was intercalated between the proximal row and the 4 distal tarsals. 
In Aelurognathus microdon (fig. 10, a) I have found 3 proximal 
elements and 4 distals; the fibula articulates with a large flattened 
fibulare which is, antero-postaxially, thin and plate-like; intercalated 
between the ends of the tibia and fibula a remarkably shaped bone, 
carrying three facets, is present; this I consider to be an intermedium, 
although it is possible that it is a composite element consisting of a 
fused intermedium and tibiale; distal to the end of the tibia there lies 
a pebble-like bone which I have identified as the tibiale, although it 
must be admitted that it does not actually lie articulated to the tibia 
and may possibly have fitted in between the intermedium and the 
distal tarsals and would then have the topographical relations of a 
centrale (navicular). There are 4 distals, but the shape of the 4th 
clearly indicates that it is a composite bone consisting of a conjoined 
4th and 5th distal tarsal. In Hipposaurus boonstrar (fig. 11), which 
comes from the Tapinocephalus zone, there are only two proximal 
tarsal elements; articulating with the fibula there is a large element 
of very distinctive shape and having a strong heel-like spur; articulat- 
ing with the tibia there is a composite element, dorsally carrying two 
rounded bosses; this element I identify as a conjoined intermedium 
and tibiale, although I admit the possibility that it may be composed 
of the intermedium and centrale (navicular). The position I have 
taken up here gives rise to some points of interest—in the geologically 
older Hipposaurus there are only two elements in the proximal row, 
whereas in the younger Aelurognathus there are three; such a position 
would be contrary to evolutionary theory. If, on the other hand, 
VOL. XXXI, PART 2. Ny 


170 Annals of the South African Museum. 


the element identified as the tibiale in Aelurognathus is really the 
centrale (navicular), the same objection could be raised since in 
Hipposaurus there is no centrale (navicular). I see no way of clearing 
up this difficulty, and am content to await some further discoveries of 
Gorgonopsian tarsi before deciding one way or the other. 

In respect of the distal tarsals of Hipposaurus there are also some 
interesting possibilities. As preserved there are 4 elements; the 4th 
element, as 1s generally the case in Therapsids, is large and obviously 
composite, and includes the 4th and 5th distal tarsals, but as it also 
stretches proximally to the 3rd distal it appears probable that it may 
also include a centrale (the 2nd); contrary to the general condition 
in Therapsids the Ist distal is a large bone, and as it stretches proximal 
to the 2nd distal tarsal it may be fused to a centrale (the navicular 2%). 
According to this view it would appear that in the tarsus of Hippo- 
saurus we have some evidence which tends to show that in the 
ancestors of the Gorgonopsia the tarsus was composed of three 
proximal elements (tibiale, fibulare, and intermedium), two centrals, 
and 5 distal tarsals. 

In no Gorgonopsian are all the digits preserved; in Lycaenops the 
Ist digit has 2 segments; in Hipposaurus the 3rd digit has 4 segments, 
and the 4th also has 4 segments. Broom’s reconstruction of the pes 
of Lycaenops, with the phalangeal formula 2, 3, 4, 5, 3, is, in view of 
the condition in Hipposaurus, probably incorrect. The Gorgonopsian 
phalangeal formula must, on the facts at our disposal, be taken as 2, 
3, 4,4, 3. The 2nd phalanx of the 3rd digit of Hipposaurus is much 
reduced, and it is probable that in the geologically younger form it is 
lost, and the phalangeal formula would then be 2, 3, 3, 4, 3. 

The chief measurements are: 


[TABLE 


A Contribution to the Morphology of the Gorgonopsia. 171 


Hippo- Aeluro- Scylacops 
SQUTUS gnathus capensis. 
boonstrai. microdon. 


S.A.M. Cat. | S.A.M. Cat. | S.A.M. Cat. 
No. 8950. No. 9344. No. 10188. 


Length : 
Kemur  . : ; J 167 2 176 2 
Tibia 5 : 4 : 159 136 95 
Fibula ; ; : f 162 sm 95 
Ath metatarsal . ; : 4] 34 2 
| Width : 
Proximal end of femur ; 30 2 32 2 
Distal end of femur . ; 33 34 Q 
Across the trochanter ; 30 2 31 UY 
Proximal end of tibia A 26 4] 25 
Distal end of tibia . : 20 Path 21 
Proximal end of fibula : RY DaTh 9 
Distal end of fibula . / an 25 12 


Comparative Morphological Discussion. 


Structurally the vertebral column of the Gorgonopsia stands on a 
higher developmental niveau than that of the Cotylosaurs in that it 
no longer possesses the heavy neural arches with low and massive 
neural spines which are so characteristic of these primitive reptiles. 
The deeply amphicoelous nature of the centra is, however, primitive, 
and in the structure of the cervical vertebra many temnospondylous 
characters are retained. This is, however, applicable to most 
Therapsids. The Gorgonopsian atlantal arches differ from those of 
Mammals, Cynodonts, and Titanosuchids in remaining separated, and 
agree in this character with the Tapinocephalids and Anomodonts. 
The presence of a dichocephalous atlantal rib is of great interest. No 
extreme adaptive specialisations such as the development of zygo- 
sphenes, zygantra, hyposphenes, hypantra, exapophyses have been 
observed in the Gorgonopsia. The apparent presence of distinct 
lumbar vertebrae is of interest. Distinct lumbar vertebrae are known 
in some Cotylosaurs (Limnoscelis, Labidosaurus), Pelycosaurs (Sphena- 
codon) and Cynodonts (Thrinaxodon). The presence of 27 to 28 
presacrals is typical of Therapsids, where this number varies between 
23 to 28; and 3 sacrals are a good average for Therapsids where this 
number varies between 2 to 7. 


172 Annals of the South African Museum. 


The shoulder-girdle of the Gorgonopsia is typical of primitive 
reptiles in containing scapula, coracoid, procoracoid, clavicle, inter- 
clavicle, and cleithrum; but in many respects shows advanced 
characters—the cleithrum is rudimentary and the glenoid is a fairly 
simple posteriorly directed fossa with only a slight indication of its 
ancestral “‘screw-shape.’’ No acromion is, however, developed as in 
Anomodonts and Cynodonts. In the geologically older Hipposaurus, 
the great length of the coracoidal plate is more Deinocephalian-like 
(Jonkeria) than in the younger Aelurognathus and Arctognathoides. A 
well-ossified sternum is present as in Anomodonts and Therocephalians. 

The great length of the pubis in Hipposaurus and Lycaenops is 
strikingly different from the condition in other Therapsids where the 
pubis is invariably much shorter than the ischium. In the Deino- 
cephalians and Anomodonts there is a single opening, the pubic 
foramen, whereas in Therocephalians and Cynodonts there is a large 
pubo-ischiadic fenestra; although not well preserved in the known 
Gorgonopsia, it appears highly probable that a pubo-ischiadic fenestra 
was developed and was probably confluent with its fellow of the other 
side. 

The humerus is a much slenderer bone with less expanded ends than 
in the Deinocephalians, Anomodonts, and Cynodonts; the delto- 
pectoral crest is also less developed than in these forms. In some 
Therocephalians, however, the humerus is even more slender and the 
shaft longer than in the Gorgonopsia. 

The epipodial of the Gorgonopsia is long and slender as in the 
Therocephalians, whereas in the Deinocephalians it is extremely short. 

The Gorgonopsian carpus contains three bones in the proximal row, 
2 centrals, and 4 or 5 distals, as in the Cynodonts; the phalangeal 
formula is 2%, 3%, 3, 3, 3% in Lycaenodontoides, in Aelurognathus 
2%, 3, 4, 4, 3, whereas in the Anomodonts it is 2, 3, 3, 3, 2 and in 
Ceedonts PAS 8) Seay os 

The femur of the Gorgonopsia is a long slender bone with a long shaft 
and unexpanded ends, whereas in the Deinocephalians (except in some 
undescribed material collected by me at Merweville) and Anomodonts 
it is short with widely expanded ends; in the Therocephalians and 
Cynodonts the femur is as in the Gorgonopsia; in the former, however, 
the internal trochanter is much stronger than in the Gorgonopsia. 

In the Gorgonopsian tarsus there are 2 to 3 separate elements in 
the proximal row; in the middle row 0 to 1; in the distal4; The 
phalangeal formula of the Gorgonopsia is 2, 3?, 4, 4, 3%. 


21. 


22. 


A Contribution to the Morphology of the Gorgonopsia. 173 


BIBLIOGRAPHY. 


. Broom, R., 1910.—‘‘ A Comparison of the Permian Reptiles of North America 


with those of South Africa,’’ Bull. Amer. Mus. Nat. Hist., vol. xxviii. 


. Broom, R., 1910.—* Observations on some Specimens of South African Fossil 


Reptiles Preserved in the British Museum,” Trans. Roy. Soc. 8. Afr., 
vol. ii. 


. Broom, R., 1911.—‘‘On some New South African Permian Reptiles,”’ P.Z.S. 
. Broom, BR., 1912.—‘‘On some New Fossil Reptiles from the Permian and 


Triassic Beds of South Africa,”’ P.Z.S. 


: Broom, R., 1913.—“On Four New Fossil Reptiles from the Beaufort Series, 


South Africa,’ Rec. Albany Mus., vol. ii. 


. Broom, R., 1913.—“‘ On the Origin of the Mammalian Digital Formula,” Anat. 


Anzeiger, Band 43. 


. Broom, R., 1913.—‘‘On the Gorgonopsia, a Sub-order of the Mammal-like 


Reptiles,” P.Z.S. 


. Broom, R., 1913.—“On some New Carnivorous Reptiles,’ Bull. Amer. Mus. 


Nat. Hist., vol. xxxii. 


. Broom, R., 1913.—‘‘On a Nearly Perfect Skull of a New Species of the 


Gorgonopsia,”” Ann. S. Afr. Mus., vol. xii. 


. Broom, R., 1914.—Croonian Lecture “On the Origin of Mammals,” Phil. 


Trans. Roy. Soc., B, vol. ccvi. 


. Broom, R., 1915.—‘‘On some New Carnivorous Therapsids in the Collection 


of the British Museum,” P.Z.S. 


. Broom, R., 1915.—‘‘Catalogue of Types and Figured Specimens of Fossil 


Vertebrates in the American Museum of Natural History. II. Permian, 
Triassic, and Jurassic Reptiles of South Africa,’ Bull. Amer. Mus. Nat. 
Hist., vol. xxv. 


. Broom, R., 1921.—‘‘On the Reptilian Tarsus,” P.Z.S. 
. Broom, R., 1925.—‘‘On some Carnivorous Therapsids,’’ Rec. Albany Mus., 


vol. ili. 


. Broom, R., 1930.—‘‘On the Structure of the Mammal-like Reptiles of the 


Sub-order Gorgonopsia,’”’ Phil. Trans. Roy. Soc., B, vol. cexviii. 


. Broom, R., 1931.—‘‘ Notices of some New Genera and Species of Karroo Fossil 


Reptiles,’ Rec. Albany Mus., vol. iv. 


. Broom, R., 1932.—‘‘The Mammal-like Reptiles of South Africa,’ Witherby, 


London. 


. Broom, R., and Havenron, S. H., 1913.—‘‘On a New Species of Scymno- 


gnathus (S. tigriceps),” Ann. 8. Afr. Mus., vol. xii. 


. Haveuton, S. H., 1915.—‘‘On some New Gorgonopsians,”’ Ann. 8. Afr. Mus., 


vol. xii. 


. Haventon, S. H., 1918.—‘‘Some New Carnivorous Therapsida, with Notes 


upon the Brain-case in Certain Species,” Ann. 8. Afr. Mus., vol. xi. 
Haveuton, 8. H., 1926.—‘‘On Karroo Vertebrates from Nyassaland,” Trans. 
Geol. Soc. 8. Afr., vol. xxix. 
Haveuton, S. H., 1929.—‘‘On some New Therapsid Genera,” Ann. S. Afr. 
Mus., vol. xxviii. 


174 Annals of the South African Museum. 


23. Haveutoy, S. H., 1932.—‘ On a Collection of Karroo Vertebrates from Tangan- 
yika Territory,” Q.J.G.S., vol. Ixxxviii. 

24. LYDEKKER, R., 1890.—Cat. Foss. Rept. Amphibia, British Mus., vol. iv. 

25. Norosa, F. von, 1923.—“‘ Die Familien der Reptilien,” Fortschr. Geol. Pal., ii. 

26. Norcesa, F. von, 1928.—‘‘ Palaeontological Notes on Reptiles,’’ Geol. Hun- 
garica, 1. 

27. OwEN, R., 1860.—‘‘On some Reptilian Fossils from South Africa,” Q.J.G.S., 
vol. xvi. 

28. OweEn, R., 1876.—Cat. Foss. Rept. S. Afr. 

29. OwEN, R., 1876.—‘‘ Evidences of Theriodonts in Permian Deposits elsewhere 
than in South Africa,” Q.J.G.S., vol. xxxii. 

30. OweEn, R., 1881.—“‘On the Order Theriodontia, with a Description of a New 
Genus and Species (Aelurosaurus felinus),” Q.J.G.S., vol. xxxvii. 

31. PRAVOSLAVLEV, P. A., 1927.—“III, IV. Gorgonopsidae,” Akad. Nauk. 

32. PRAVOSLAVLEV, P. A., 1927.—‘“‘Relation of the Gorgonopsids of Northern 
Russia to Mammals,”’ Ann. Soc. Pal. Russe, vol. i. 

33. SEELEY, H. G., 1895.—‘‘Researches on the Structure, Organisation, and 
Classification of the Fossil Reptilia. Part IX, Section I. On the Thero- 
suchia,”’ Phil. Trans. Roy. Soc., B, vol. clxxxv. 

34. Watson, D. M.S., 1912.—‘‘On some Reptilian Lower Jaws,”’ Ann. Mag. Nat. 
Hist., Ser. 8, vol. x. 

35. Watson, D. M. S., 1913.—‘‘On some Features of the Structure of the Thero- 
cephalian Skull,” Ann. Mag. Nat. Hist., Ser. 8, vol. xi. 

36. Watson, D. M. S., 1914.—‘* Notes on some Carnivorous Therapsids,” P.Z.S. 

37. Watson, D. M. S., 1921.—‘‘The Bases of Classification of the Theriodontia,” 

PAS: 


(175 ) 


6. Additions to our Knowledge of the South African Gorgonopsia, pre- 
served in the British Museum (Natural History).—By Lizuwe D. 
Boonstra, D.Sc., Palaeontologist to the South African Museum 
and Queen Victoria Scholar of the University of Stellenbosch. 


(With 18 Text-figures.) 


THERE is preserved in the British Museum (Natural History) a 
small, though historically important collection of South African 
Mammal-like Reptiles of the sub-order Gorgonopsia. In all there 
are thirty specimens. These have been studied by Owen, Lydekker, 
Seeley, Broom, and Watson, and have been referred to fifteen genera 
and sixteen species. As will become apparent in the sequel, some 
of these so-called species cannot be considered to have been founded 
on valid characters. There are included, one specimen which 
apparently comes from the Tapinocephalus zone, six species from the 
Endothiodon zone, and nine species from the Cvrstecephalus zone. 
Considered as a whole, this collection consists of very indifferently 
preserved material. Quite a number are of a very fragmentary and 
incomplete nature, and some show practically no structure at all; 
even the best preserved specimens leave much to be desired. A 
considerable amount of preparation has been necessary, and the 
following descriptions are, I believe, as full as the nature of the 
material warrants. 


A. Forms from the Tapinocephalus zone. 
Cymscodon lydekker1, Broom. 


The type-specimen is the anterior portion of a small right dentary. 
It was found on the farm Palmietfontein by T. Bain in 1878; but 
as there are many farms in the Karroo with this name it is not certain 
that the specimen represents a form from the Tapinocephalus zone. 
It was found in association with some remains of a small Dicynodont, 
and was referred by Lydekker to Cynosuchus suppostus. Broom in 
1915 showed that the dentary was apparently that of a Gorgonopsian. 
The roots of three incisors, one canine, and four small molars are 
preserved. Broom was of the opinion that in life four incisors were 
present; I can find no evidence for this supposition; as preserved, 


176 Annals of the South African Museum. 


the first incisor is practically on the anterior edge of the bone and 
there is no room for an additional tooth. There is no diastema 
anterior to the canine, which has an oval root and lies lateral to the 
last incisor. Posterior to the canine, there is a diastema of 7 mm.; 
the three incisors occupy 7 mm.; the canine measures 6 by 3 mm., 
and the four molars measure9 mm. The mentum is deep and square; 
the depth being 19 mm., and the width 20 mm. 

At the present time five types of Gorgonopsia from the Tapino- 
cephalus zone are known, viz. Hoarctops, Eriphostoma, Galesuchus, 
Hipposaurus, and Scylacognathus. Of these Hriphostoma is the 
only one to which the fragment of Cyniscodon can be compared. 
The nature of the mentum is very similar, but the dentition is quite 
different. Broom compared it to Aelurosaurus, but in this form the 
mentum is sloping, whereas in Cyniscodon it is upright; in addition, 
there is a considerable difference in the nature of the teeth. The 
relations of this fragment thus remain unknown. 

As this fragment throws practically no light on the nature of the 
Gorgonopsians of the Tapinocephalus zone, it seems a pity that a 
new genus should have been created for such an unimportant frag- 
ment with practically no diagnostic characters. An additional 
name does not increase our knowledge. 

Type, B.M.N.H., 49404, Palmietfontein, Beaufort West?, Cape 
Province. 


B. Forms from the Endothiodon zone. 


From this zone there are in the collection six forms, viz. Arctops, 
Aelurosaurus, Aelurosauroides, Gorgonops, Leptotrachelus, and 
Scymnognathus. Most of these are represented by fairly complete 
skulls, from which a number of structural details has been determined. 


Arctops willistoni, Watson. 
(Figs. 1-3.) 


This type was collected by A. G. Bain in 1857 at Howse Post, near 
Fort Beaufort, which locality Watson considered to be in the 
Endothiodon zone; the nature of the matrix suggests the older 
Tapinocephalus zone. In either case, Arctops is one of the earlier 
Gorgonopsians. It consists of the posterior two-thirds of a skull, 
from which the quadrates and quadratojugals have been lost. Watson 
in 1914 described and figured the occiput and part of the brain-case, 
with which account I am in agreement. In 1921 figures and a 


South African Gorgonopsia preserved in the British Museum. 177 


description of the dorsal, lateral, and palatal surfaces were published. 
I have since etched all these surfaces with dilute hydrochloric acid 


‘ ‘ 


ey _ Se 
’ 
y ay \ EN 
Ye 6 't- ‘-/ 0 yy 
‘ot te J ewan 
/ ae | ! i] ‘ 
1 ' i) J \ 
/ i ty \ 
/ a / 


Oe ie Pes ROe SOnvtcOc Techs 


Fig. 1.—Arctops willistoni. Type, B.M.N.H., R4099. Dorsal view of the 


skull. x 4. 
B.Oc. =basioccipital. P.Fr. =postfrontal. 
Ex.Oc. =exoccipital. P.Oc. =paroccipital. 
I.Pa. =interparietal. P.Orb. = postorbital. 
13Tes = frontal. Pr.Fr. =prefrontal. 
Ju. =jugal. Pr.Pa. = preparietal. 
La. =lacrimal. S.0c. =supraoccipital. 
Mx. =mazxilla. Sq. =squamosal. 
Na. =nasal. Tab. =tabular. 


Pa. §=parietal. 


and developed the brain-case, so that the skull now shows a number 
of points not determined by Watson. This account is thus purely 
supplementary. 


178 Annals of the South African Museum. 


On the dorsal surface the radiation of the bone-fibres assists 
materially in determining the limits of the various bones, which are 
best understood by referring to fig. 1. The short cruciform frontal 
deserves notice. 

On the ventral surface I was able to determine that the palatines 
meet in the median line, and thus on the palatal surface the pterygoids 


~ iy ce SS cess eeetan Aue ais S as Se 
41a n mh te Nah BN dass y vias el gt). 


Fie. 2.—Arctops willistoni. Type. Ventral view of the skull, modified after 
Watson. x 4. 


B.Sph.  =basisphenoid. Pter. =pterygoid. 
Ket.Pter. =ectopterygoid. Pr.Ot. = pro-otic. 
Pal. = palatine. Pr.V. =prevomer. 


are prevented from meeting the prevomers; on either side of the 
median line there is a low ridge; enclosed between these ridges is a 
median groove; just behind the highest part of this ridge the transverse 
suture, between the anterior pterygoid ramus and the palatines, is 
clearly seen as a convoluted line lying anterior to a dark line of 
matrix in a crack, which runs parallel to it and with which it must 
not be confused; in a parasagittal section the pterygo-palatine 
suture can also be seen; the suture then continues in lateral direction 
to meet the ectopterygoidal suture (fig. 2). The ectopterygoid 
descends to form the anterior corner of the transverse pterygoidal 


ren 


South African Gorgonopsia preserved in the British Museum. 179 


_ process, which is much shallower and situated somewhat further 
posteriorly than in later forms, but it has shifted somewhat forward 
from the primitive condition. The junction of the prevomers and 
palatines is not clearly shown, but its nature is indicated in broken 
lines in the figure. No teeth can be seen on the median palatal 
ridges, but in analogy with other known forms it is certain that 
they were present on the palatinal portion of the ridge, but would 
appear to have been absent on the pterygoidal part. In a longi- 


Fie. 3.—Arctops willistont. Type. Lateral view of the right side of the 
brain-case. x 4. 


B.P.P. =basipterygoid process. Pit. =pituitary fossa. 
Ep.Pter. =epipterygoid. Sph.Ethm. =sphenethmoid. 
Fen.Ov. =fenestra ovalis. Ven. =venous fossa. 


P.Sph. =parasphenoid. 
Roman numerals refer to the foramina for the cranial nerves. 


tudinal section, it is seen that the anterior pterygoidal ramus carries 
a high dorsal keel. 

The palate of Arctops is thus seen to agree in the essential points 
of structure with all the forms in which the palate has hitherto been 
studied, viz. Scylacops, Aelurognathus, Arctognathus, Lycosaurus, 
Cynariodes, Lycaenodon, Lycaenodontoides, and Arctognathordes, and, 
as I hope to show later in this paper, with Gorgonops, Aelurosaurus, 
Aelurosauroides, and Scymnognathus. In all these forms there is 
no evidence whatsoever of a median unpaired bone in the anterior 
part of the palate. 

The basicranial region is not very well shown; the basisphenoidal 


180 Annals of the South African Museum. 


tubera are present as high narrow ridges with a broad deep groove 
in between; posteriorly, it rests on ridge-like tubera of the basi- 
occipital, and it forms the anterior border of the fenestra ovalis; 
anteriorly, the limits of the basisphenoidal portion of the median 
keel and the basipterygoidal processes cannot be determined. Al- 
though I have gone to considerable trouble to expose the lateral 
surface of the brain-case on the right side, the result is rather dis- 
appointing. The posterior part of the quadrate ramus of the 
pterygoid is lost, and portions only of the epipterygoid, which rested 
on it, can be indistinctly seen; it has also been impossible to expose 
the sphenethmoidal part of the brain-case satisfactorily. Pos- 
teriorly, the broad anterior end of the basioccipital and the widely 
diverging basisphenoidal tubera produce a roomy floor to the brain- 
case; the skull is also high in this region, but due to the shortness of 
the parietal, the frontal with the sphenethmoid attached to its 
ventral surface lies far back, thus producing an extremely short 
hind brain. The pro-otic stands on the basisphenoid and les against 
the anterior face of the paroccipital in the usual manner; it 1s, 
however, an extremely small bone, which stretches very little in 
anterior direction, and dorsally extends for less than half the height 
of the occiput. It would thus appear that in Arctops a considerable 
part of the otic region did not ossify. In Leptotrachelus and Scymno- 
gnathus the pro-otic is higher and longer, and, in the forms from the 
Crstecephalus zone (e.g. Cynariops), it is very much higher and longer. 
The lateral opening into the pituitary fossa, the notch for the fifth 
nerve, and the venous notch above it are situated far back and low 
down; the latter two openings have no ossified anterior borders as 
have the later forms. In sagittal section it is seen that the spur of 
the basisphenoid, forming the posterior border of the pituitary fossa, 
is well developed. The fenestra ovalis is large and irregular, and is 
on the level of the basioccipital condyle. Posteriorly, neither the 
supraoccipital nor the interparietal send flanges to contribute to the 
walls of the brain-case, as they do in later forms. In the median 
line the occiput is extremely thin. The parietal sends down two 
lateral flanges, with which the rod-like epipterygoids presumably 
articulate. Lying ventral to the frontals, parts of the sphenethmoidal 
portion of the brain-case can be seen. Unfortunately the epiptery- 
goid is not preserved; if it were rod-like, as in all known Gorgon- 
opsians, the sides of the middle portion of the brain-case would be 
largely unenclosed by bone, 2.e. the cavum epiptericum is not yet 
included within the cranial cavity. 


South African Gorgonopsia preserved in the British Museum. 18] 


The main morphological features are: 

Primitive. Square section of the snout; laterally directed orbits; 
very small lateral temporal openings; sides straight; Deeie central 
large; strong paroccipital; fenestrae ovales far apart; basisphenoid 
massive, but with deep edge-like tubera; pro-otic small; pineal 
foramen situated far posteriorly; piconbital depression; frontal 
forms large part of the orbital border; little ossification of the side- 
walls of the brain-case; wide parietal region. 

Advanced. Pterygoid flanges not far posteriorly; skull slightly 
wider than high; flat laterally-placed basipterygoid processes; 
maxilla apparently fairly deep; basisphenoidal tubera edge-like; 
occiput fairly upright, but not concave. 

From this summary it is evident that Arctops, although advanced 
beyond the Pelycosaurian and Deinocephalian stage, is one of the 
more primitive Gorgonopsians. 

Chief measurements : 


Pineal foramen to edge of occipital plate . 9 mm. 
Width across the squamosals . : ed NED. nek 
Interorbital width : : ; G4 ee 
Intertemporal width : ; Be OO wis, 
Width across lateral ae fauees SOON i, 
Height of snout. : : : BO". 
Height of occiput . ; : ; SOY 


Type, B.M.N.H., R4099, Howse Post, Fort Beaufort, Cape Province. 


Gorgonops torvus, Owen. 


(Fig. 4.) 

This is the type species, and was found by A. G. Bain in 1853 at 
Mildenhalls, near Fort Beaufort. This locality probably is in the 
Endothiodon zone; another specimen, in the American Museum, comes 
from an undoubted Hndothiodon zone locality, and we may thus 
consider Gorgonops to be a form from that zone. The type is a good 
skull which lacks the temporal arches and the mandibles; the occiput 
and posterior half of the ventral surface are not clearly shown, and in 
general the surface has been somewhat stripped by former developers. 

Owen’s lithographs are beautiful illustrations of the outward 
appearance of the skull as preserved. 

Watson’s account (1921) of the dorsal and lateral surfaces is correct; 
anterior to the pineal foramen, a large preparietal, which was not 


1382) Annals of the South African Museum. 


figured by Watson (fig. 4), is present; his interpretation of the rela- 
tions of the septomaxilla, maxilla, and the premaxilla is correct. 

With regard to the palate, I am in agreement with Watson as to his 
interpretation of the relations of the premaxilla and the prevomers, 
and also as to the posterior intercalation of the prevomers; but my 
interpretation of the relations 
of the palatines, pterygoids, 
and ectopterygoids is totally 
different. The posterior slip- 
like prolongation of the pre- 
vomers is clasped in the median 
line by the two palatines lying 
laterally; from the posterior 
tip of the prevomers a median 
suture proceeds in_ posterior 
direction for a short distance; 
it then bifurcates and each 
branch continues in_ postero- 
lateral direction, crossing the 
ridge, that borders the median 
groove at its lowest part, to 
meet the ectopterygoidal suture. 
According to this interpretation, 
Fic. 4.—Gorgonops torvus. Type, B.M.N.H., gine (DEES SCLLS LTE aki 

R1647. Ventral view of the skull, excluded from the posterior 

modified after Watson. x 3. borders of the internal nares; 

Pr.Mx. =premaxilla. whereas, according to Watson, 

the pterygoids stretch far for- 

ward and are intercalated between the palatines, which do then not 

meet in the median line. The limits of the ectopterygoids are not 

clearly visible, but I am inclined to think that they descend further 
down the pterygoid flanges than figured by Watson. 

The ridges bounding the median palatal groove are not well pre- 
served; but although no teeth are visible, in life they were probably 
present on both the pterygoidal and palatinal portions of these ridges. 
Watson has maintained that the roof of the posterior part of the 
median groove is formed by a distinct element—the vomer—which is, 
of course, quite distinct from the prevomer. In Gorgonops there is no 
evidence of any posterior limit to this part of the roof of the median 
vault, which would definitely separate it from the rest of the ptery- 
goid. My interpretation of the palate of Gorgonops is thus in agree- 


South African Gorgonopsia preserved in the British Museum. 183 


ment with the condition found in the nine Gorgonopsian genera 
mentioned above. 

The main morphological features of Gorgonops are: 

Primitive. Square section of the snout; laterally directed orbits; 
slightly sloping occiput; tooth row straight and step in alveolar 
border; very small lateral temporal openings; basioccipital large; 
posterior position of the pineal foramen; preorbital depression; 
frontals form large part of orbital border; large postfrontals. 

Advanced. Pterygoid flanges not far posteriorly; slightly cupped 
occiput; sides of skull not straight; snout wider than high; maxilla 
deep; flat basipterygoid processes laterally placed. 

As far as the skull of Gorgonops is known, it is quite as primitive as 
Arctops, but as the region of the nostril is absent in the latter, and the 
brain-case is unknown in the former, further discussion would not be 
profitable. 

Chief measurements: 


Length from basioccipital to premaxilla ; oy) 21 Ommama: 
Length from premaxilla to pineal foramen . LiON 8 
Length from pineal foramen to edge of occipital plate. La; 
Length from premaxilla to front of orbit. : ; ATI sate 
Width across squamosals_ . . i : vy Ore. 
Interorbital width . ; : : : : 60 ,, 
Intertemporal width : ‘ : Ge 5 
Width across the canines . : : : : : GOe 
Width across pterygoid ie ; : (Cae 
Height of snout ’ : BOK 
Height of occiput. AD 3 
Diastema : ; f : ; : . : 1b 
Incisor series. 34, 


Type, B.M.N.H., R1647, Mildenhalls, Fort Beaufort, Cape Province. 


Scymnognathus whaitsi, Broom. 
(Fig. 5.) 

There are in the collection three specimens of this species, which 
have been described by Watson. All come from the Hndothiodon zone 
of Beaufort West. 

In R4052 it is possible to determine from the weathered fragments 
the nature of the snout and the dental formula (i. 5, c. 1, m. 4 or 5), as 
fully described by Watson. The material is, however, not sufficiently 
well preserved to enable one to state with certainty what the relations 


184 Annals of the South African Museum. 


of the prevomers, palatines, and pterygoids are. Watson’s figure of 
this part is based on inconclusive evidence, and I have no doubt that 
better preserved material will show that, in this region, Scymnognathus 
agrees with other Gorgonopsians, viz. that the palatines meet in the 
median line, that the pterygoids do not meet the prevomers, and that 
there is no unpaired median bone. 

An imperfect weathered axis is the basis on which this element was 


Po. 


Vid. B. Spl. HT. 
Fic. 5.—Scymnognathus whaitsi. B.M.N.H., R4053. Lateral view of the 


left side of the brain-case, modified after Watson. x 2. 
Vid. = Vidian foramen. 


figured by Watson. The anterior cervicals are much better shown in 
Hipposaurus, Aelurognathus, and Arctognathoides recently described 
by me, and these show that Watson was correct in his restoration. 

Watson’s figures of the sectioned snout (49369) are correct. In 
fig. 12, No. 7, the inner pair of strips labelled “pterygoid” are open 
to discussion. There is no doubt as to their presence as separate 
strips of bone; what must be decided is whether they represent the 
anterior extremities of the pterygoids or not. If they do, it means 
that the palatines do not meet in the median line and that the ptery- 
goids meet the prevomers, which would be contrary to the condition 
in at least nine other Gorgonopsia. I believe that the explanation is 
that they represent a dorsal pterygoidal girder which is not exposed 
in ventral view. 


South African Gorgonopsia preserved in the British Museum. 185 


In R4053 the back portion of the skull is fairly well preserved; the 
proatlas and the axial centrum are well shown in situ; and the 
structure of the occiput and the greater part of the brain-case can be 
determined. From the material in the collection I have been 
fortunate enough to develop the brain-case in three different genera— 
Cynariops, “* Lycaenodon,” and Arctognathus—and comparing these 
to the figure of the brain-case of Scymnognathus published by Watson, 
some very disturbing differences were noticed. Re-examining the 
specimen in the light of this new knowledge, an interpretation different 
to that of Watson was arrived at. In the figure it is shown that the 
pro-otic is a much larger bone than Watson thought; its ventral limit 
is marked by a suture, which runs from behind the basisphenoidal 
tubera to the edge of the lateral opening into the pituitary fossa; 
dorsally, it extends right up to the parietal and interparietal and forms 
the ventral and dorsal borders of the notch (or foramen ?) for the fifth 
nerve, and the ventral border of the venous fossa; it is this dorsal part 
that Watson, as I believe in error, considered to be supraoccipital. 
The foramen for the sixth nerve is extraordinarily large, and lies 
ventro-posteriorly of the notch for the fifth nerve; it has a depression 
below it. The foramen for the seventh nerve lies in advance of the 
fenestra ovalis, and below it is a depression for the geniculate ganglion. 
In the specimen the basipterygoid process has been forced upwards, 
and in the figure its normal horizontal position is restored; behind 
this process lies an opening—the Vidian foramen. 

The chief morphological features are: 

Primitive. lLaterally directed orbits; sloping occiput; slender 
epipterygoid; large basioccipital; fenestrae ovales far apart; basi- 
sphenoidal tubera fairly large, but edge-like; step in maxillary 
border. 

Advanced. Snout somewhat broader than high and not square in 
section; pterygoid flange has shifted forward; basipterygoid process 
laterally situated; parietal enters brain-case; pro-otic fairly large; 
pineal foramen not posteriorly situated; practically no preorbital 
depression; frontal forms only small part of orbital border; maxilla 
deep; paroccipital not massive; basisphenoidal tubera situated more 
anteriorly than in Arctops; squamosals widened, and thus increase in 
size of temporal openings; occiput concave. 

A consideration of all these characters tends to show that 
Scymnognathus, although retaining some primitive characters in 
common with Arctops and Gorgonops, does show a definite advance 
over them. 

VOL. XXXI, PART 2. 18 


186 Annals of the South African Museum. 


Scymnognathus sps. 


In the collection is a specimen found by Professor D. M. 8. Watson 
in 1915; he did not think it worth describing, but Broom, in a paper; 
which consists of a number of specific descriptions based on absolutely 
impossible material, created a new species—S. parvus. The type isa 
weathered skull, very badly crushed; nothing can be made out of the 
palate, occiput, or lateral surface; on the dorsal surface a few sutures 
are visible. The dental formula is as for the genus—i. 5, c.1, m.4; the 
posterior edges of the incisors and canines have very definite fine 
serrations, whereas, on the last molar, which alone is fairly well 
preserved, there are some indefinite grooves which may be serrations. 

I do not think that the specimen is sufficiently well preserved to 
warrant the creation of a new species; the few characters that can be 
determined are sufficient only to identify the genus, and there are 
no valid characters to differentiate it from the other species of 
Scymnognathus. 

B.M.N.H., R4139, Kuilspoort, Beaufort West, Cape Province. 


There is another specimen consisting of a fragment of a left maxilla, 
which shows the dental formula—i. 4 or 5, c. 1, m. 3, and has a distinct 
step anterior to the canine. This has, I believe, been correctly 
identified by Broom as Scymnognathus sp. 

B.M.N.H., R3611, Karroo. 


Leptotrachelus eupachygnathus, Watson. 
(Figs. 6-7.) 


Although somewhat distorted, the skull shows a considerable 
portion of its structure, and this has been fully described by Watson. 
The dental formula, which is the same as in Scymnognathus whaitsi, is 
i.52,c.1,m.4 or 5. There is no evidence as to the nature of the 
crowns, and no definite evidence of a step in the alveolar border as 
Watson’s figure would imply. I have refigured the dorsal aspect of 
the skull, for, as Watson has admitted, the snout as reconstructed by 
him is too long and narrow. The position of the pineal foramen has 
also been determined. 

I have refigured the brain-case in order to indicate the position of 
the basisphenoidal-pro-otic suture, the depression which apparently 
leads into a venous fossa, and the probable position of the foramen for 
the sixth nerve. Above the notch for the fifth nerve the bone has 


Fic. 6.—Leptotrachelus ewpachygnathus. Type, B.M.N.H., R4051. Restored 
dorsal aspect of the skull. x 4. 


B.Sph. Viel. VIL. Fen. Ov: 
Fic. 7.—Leptotrachelus ewpachygnathus. Type. Lateral view of the left side 
of the brain-case, modified after Watson. x 2. 
P.T.F. =posttemporal fenestra. 


188 Annals of the South African Museum. 


suffered from crushing and sun-cracking, and it can unfortunately not 
be determined to what extent the interparietal and supraoccipital 
contribute to the formation of the lateral wall of the brain-case. 

The brain-case of Leptotrachelus is thus very similar to that of 
Scymnognathus, and shows, particularly in the size of the pro-otics and 
the part played by the parietals, interparietals, and to a lesser extent, 
the supraoccipitals, a stage of development well in advance of that 
attained by the more primitive Arctops. In this form, attention was 
drawn to the fact that the pro-otic was not developed anterior to the 
notches identified as pituitary, venous, and fifth nerve openings; in 
both Leptotrachelus and Scymnognathus these notches are deeper, 1.e. 
more enclosed by the pro-otic. As will be pointed out when dealing 
with Cynariops and “‘ Lycaenodon,” some of these notches are wholly 
enclosed by bone in these forms. 

The chief morphological features are: 

Primitive. Preorbital depression; frontals forming large part of 
orbital border; apparent step in alveolar border; large postfrontal; 
large quadrate. 

Advanced. Rounded snout, fairly depressed; deeply cupped but 
upright occiput; squamosal bayed, forming fairly large temporal 
opening; curiously small basioccipital; basisphenoid far forward; 
pro-otic fairly large; deep maxilla. 

Leptotrachelus is thus very similar to Scymnognathus and stands 
on the same developmental niveau. 

Chief measurements : 


Length from premaxilla to basioccipital condyle . . 170? mm 
Length from premaxilla to pineal foramen . . 1 L20%aae 
Length from pineal foramen to edge of occipital plate : 24 ,, 
Length from premaxilla to front of orbit — . 1 x, 00 Gee 
Length of lower jaw . ; 200 ue 
Width across squamosals_ . : : , : [60.3 
Interorbital width . ; : 4 : : HO ae 
Intertemporal width . : : . 5 ; : SOs 
Height of snout : é : ; : : GD) 
Height of occiput : : : ; : é Gl aes 
Height of mentum of lower jaw . : Dil 


Type, B.M.N.H., R4051, Hans River, Beaufort West, Cape Province. 


South African. Gorgonopsia preserved in the British Museum. 189 


Aelurosaurus felinus, Owen. 
(Fig. 8, A.) 

Owen’s original figures excellently illustrate the form and general 
appearance of the skull, but he mistakenly considered it “‘ mononarial.”’ 
Lydekker rightly pointed out that in reality the internasal bar was 
lost. Owen’s dental formula—i. 5, c. 1, m.5—is correct, and Lydekker, 
in his description and figures, drew attention to the finely serrated 


A, 


Ect. Prer. 
Pher 


Fig. 8. 


A.—Aelurosaurus felinus. Type, B.M.N.H., R339. 
B.—Aelurosauroides watsoni. Gen. et sp.nov. B.M.N.H., R855. 


Ventral views of the anterior part of the palate. x. 


nature of the posterior borders of both incisors and molars. Seeley’s 
interpretation of the articulation of the lower jaw was at fault, and 
Broom (1910) showed that in Aelurosaurus this structure is typically 
as in other Gorgonopsians. Seeley correctly noticed teeth on the 
transverse pterygoidal bar, and also noticed two groups of teeth on 
the pair of palatal ridges, but failed, as did Broom and Watson after 
him, to see the transverse suture between them. This suture between 
the palatine and pterygoid is plainly visible, on the left side, as a 
dark convoluted line demarcating the two lighter coloured bones. 
From the ectopterygoidal suture, this suture runs medially and 
crosses the tooth-bearing ridge between the two sets of teeth—an 
anterior larger set on the palatine, and a set of fewer and smaller 
teeth on the pterygoid. Unfortunately the ectopterygoid is not 


190 Annals of the South African Museum. 


sufficiently exposed, but it apparently stretched for a considerable 
distance down the transverse pterygoidal flange. 

The main morphological features are: 

Primitive. Snout higher than wide; laterally directed orbits; 
slight preorbital depression; frontals form large part of orbital 
border. 

Advanced. Snout rounded; anterior position of pterygoidal 
flanges; deep maxilla; no step in alveolar border. 

The characters shown by Aelurosaurus, and also by Aelurosauroides, 
indicate that these two forms occupy a position quite distinct from 
that of the other forms from the Endothiodon zone. It would appear 
that they represent a branch stock of the Gorgonopsia, in which the 
more advanced characters of the later Gorgonopsians are, as 1t were, 
foreshadowed. 

Chief measurements : 


Length from premaxilla to pineal foramen . . 80 mimi: 
Length from premaxilla to front of orbit . Baa 
Length of lower jaw. ; : . oe 
Interorbital width . : ; 1 : . sae 
Width of snout f , Joo 
Width across lateral peeeeidal flanges ; 5 ea ee 
Height of snout ; ‘ é > eee 
Height of mentum of ips jaw. 3 ; 2° Toke 
Length of molar series. 4 . ‘. whey 
Diastema : P ; : : : : Gwe 
Length of incisor series. 1G 


Type, B.M.N.H., R339, Gouph, Beaufort West, Oats Province. 


Aelurosauroides watsoni, gen. et sp. nov. 
(RionGs 5.) 


This specimen has been regarded by both-Lydekker and Broom 
as Aelurosaurus felinus. It consists of the anterior two-thirds of 
a small skull, and lacks the lower jaw; the outer surfaces are well 
exposed, and Watson has had the palate developed. I concur in 
Watson’s description and figures, except that, in dorsal view, the 
preparietal is oval and not squarish and, moreover, agree with him 
that it is generically distinct from Aelurosaurus. In addition to 
the points enumerated by him, I have been able to determine that 
the palatines meet in the median line, and thus the pterygoids do not 
meet the prevomers. 


South African Gorgonopsia preserved in the British Museum. 191 


Aelurosauroides agrees with Aelurosaurus felinus in its dentition, 
size, and general shape, but differs in that the snout is rounder and 
less high. As Watson has already pointed out, the two forms differ 
very markedly in the relative size and shape of the palatal elements, 
although they agree in the essential morphological features. Watson’s 
point, that great care must be taken before assigning specimens of 
Gorgonopsians, which have the same outward appearance, to the 
same species, without knowing the nature of the palate, therefore 
deserves full emphasis. The differences in proportion are best 
understood by referring to the figures; the main are—the shortness 
and raised nature of the tooth-bearing ridge in Aelurosauroides; the 
relatively shorter prevomers; the great ventral extent of the ecto- 
pterygoids, and the more posterior extent of the internal nares. 

The main morphological features are: 

Primitive. Laterally directed orbits; snout slightly higher than 
broad; slight. preorbital depression; frontal apparently forms a 
large part of the orbital border. 

Advanced. Snout short and somewhat rounded in section; 
pterygoid not situated far posteriorly; tooth row slightly curved; 
sides not straight; deep maxilla; no step in alveolar border. 

Thus, although retaining some characters which are primitive, 
Aelurosauroides is more advanced than the forms from the Tapino- 
cephalus zone, and also than Arctops and Gorgonops. 

Chief measurements : 


Length from premaxilla to front of orbit . sy) (oO innrcay 
Interorbital width . ; i : : Asuka Pa 
Width of snout : : : : 4 yy Oot ee 
Height of snout : : : Roce oe 
Width of pterygoid flanges “ My OOr 5s 
Length of molar series. ates lige 
Diastema : : : : : : (Shite 
Length of incisor series. s : Spee ovals 


Type, B.M.N.H., R855, Gouph, Cape Province. 


An Unidentified Specimen. 


There is in the collection a specimen, labelled 855a, which lacks 
the external surface; but its palate can be exposed. Until this is 
done, it is not possible to state whether it must be referred to Aeluro- 
saurus or to Aelurosaurovdes. 


192 Annals of the South African Museum. 


C. Forms from the Cistecephalus zone. 

Hight species from this zone are represented, viz. Arctognathus, 
Arctosuchus, Cerdognathus, Cynariops, Cynodraco, Lycaenodon, 
Lycosaurus, and Scylacops. Careful preparation has produced two 
good palates and the lateral surfaces of three excellent brain-cases. 


Lycaenodon longiceps, Broom. 
(Fig. 9.) 


The type-specimen consists of two-thirds of a skull; the whole 
parietal, occipital, and basicranial regions are lost; very little of 
the palate is present. A short original description was given by 
Broom in 1925, and in 1930 two figures were published. The dental 
formula isi. 5, c. 1,m.6; the very long molar series, measuring 28 mm., 
is remarkable, as in the Gorgonopsia the tooth row is in general very 
short. A sagittal fracture reveals some points of structure, which 
were figured by Broom; the dorsal unperforated keel of the pre- 
vomers is very strongly developed; the sphenethmoid is a large 
element, and its shape is probably as figured by Broom; on its antero- 
lateral surface there is a large foramen for the exit of the second nerve; 
Broom’s figure clearly shows that the palatines meet in the median 
line and thus prevent the pterygoids from meeting the prevomers. 
There can be no doubt that this is the usual Gorgonopsian condition, 
and that my interpretation of the palatal aspect of Arctops, Gorgonops, 
Arctognathus, etc., is correct, and that Watson misinterpreted these 
relations. Broom’s reconstruction of the basicranial portion of the 
sagittal section, as figured, is based on another specimen, and this 
will be referred to later. The detailed structure of the anterior part 
of the palate cannot be determined. Its general shape is, however, 
of interest; although the skull is a long one, the prevomers appear 
to be relatively longer, and the palatines do not stretch so far 
anteriorly, as 1s usually the case—their anterior borders lie well 
posterior to the plane of the canines; the median groove is deep, 
but unfortunately the ridges bounding it are not shown. 

The outer surface, although thoroughly cleaned and etched, shows 
few sutures, as the skull appears to be that of an old animal with 
closed sutures. The dorsal surface, with the posterior third recon- 
structed, is shown in the accompanying figure. 

The main morphological features are: 

Primitive. High, fairly narrow snout; extraordinarily long 


South African Gorgonopsia preserved in the British Museum. 193 


straight tooth row, with a step anterior to the canine: preorbital 
depression; frontals apparently forming a large part of the orbital 
border; temporal openings apparently small, and parietal region 
probably very wide. 

Advanced. Orbits slightly anteriorly directed; sides of snout 
not upright; deep maxilla. 


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Fie. 9.—Lycaenodon longiceps. Type, B.M.N.H., R5700. Restored dorsal 
view of the skull. x 4. 

As far as the skull of Lycaenodon is known, it appears to be very 
primitive for a form from the base of the Cistecephalus zone. The 
very long molar series is unique amongst all the known Gorgonopsia, 
and, if we are correct in considering this to be a primitive character, 
then Lycaenodon is in this respect the most primitive known Gor- 
gonopsian. The general contour indicates a very broad parietal 
region, with very small laterally directed temporal openings, and if 
such is actually the case, then Lycaenodon would be less advanced in 
this region than either Arctops or Gorgonops. 


194 Annals of the South African Museum. 


Chief measurements : 


Length from premaxilla to front or orbit. . 100% mm. 
Interorbital width : y . : J Aes 
Width of snout. , . : , ; oD 
Width across lateral pterygoidal flanges . st) Boas 
Height of snout . ; : ; d : AG 
Length of molar series . J : ; : ee 
Diastema . : : 3 ; : 10, 2 
Length of incisor series . f 4 : ; ise 


Type, B.M.N.H., R5700, Biesjespoort Station, Cape Province. 


“ Lycaenodon”’ sp. 
(Fig. 10.) 


In the collection purchased from Dr. Broom, in October 1932, 
there is included an isolated brain-case, parts of which were figured 
in 1930 by Broom, and included in his reconstruction of the sagittal 
section of Lycaenodon, and it is stated that it probably belongs to 
this species, presumably on the fact that it was found at the same 
locality. The difference in the state of preservation precludes the 
assumption that it belongs to the same individual, and, as this 
particular part is not preserved in the type, a comparison is not 
possible. Until further specimens are found, there is no means of 
proving that the brain-case is that of Lycaenodon; for convenience 
of reference it will be referred to “‘ Lycaenodon”’ sp. 

The whole posterior part of the brain-case and basicranial axis is 
perfectly preserved, and I have, with great care, completely exposed 
the brain-case on the left side and removed all but the dorsal extremity 
of the epipterygoid; the anterior border of the pro-otic and the 
pituitary fossa have also been freed of matrix. This specimen, before 
adequate development, was the basis of Broom’s (1930) figure showing 
the relations of the epipterygoid, pterygoid, and the two elements 
identified as “vomer”’ and ‘“‘basisphenoid”’ respectively, and also 
of the reconstructed posterior part of his sagittal section of Lycaenodon 
longiceps. 

The basisphenoid is a fair-sized bone with edge-like tubera, which 
underlie the processes sent downwards by the basioccipital, and 
which form the anterior border to the large irregular fenestra ovalis. 
From the dorso-anterior corner of the fenestra a slightly digitating 
suture runs anteriorly, in a plane practically horizontal, to enter the 


South African Gorgonopsia preserved in the British Museum. 195 


lateral border of the pituitary fossa. From the tubera the basi- 
sphenoid extends, in anterior direction, in the form of a vertical 
sheet of bone, which laterally carries the horizontal basipterygoid 
processes; on the ventral surface this produces the median keel; 
dorsally, it forms the anterior border of the pituitary fossa. Here 
it is broken off, but, in life, it apparently stretched forwards and 
upwards as a median septum. In Arctognathus curvimola this 


tr i | 


Fic. 10.—‘‘ Lycaenodon.” B.M.N.H., R5746. Lateral view of the left side 
of the brain-case. x 4. 


structure is better shown, and this will be discussed further when 
describing that form. It is evident that this extent of bone cannot 
all be basisphenoid; the anterior part of the ventral keel, and the 
antero-dorsal vertical sheet of bone must, for its greater part, be the 
parasphenoid. The limits of the parasphenoid cannot, however, 
be determined; the parasphenoid—a membrane bone—appears to 
have become intimately connected with the basisphenoid, which 1s, 
of course, preformed in cartilage. 

As indicated in the figure, the flattened, horizontally situated 
basipterygoid processes of the basisphenoid are clasped on their 
dorsal and ventral surfaces by the posterior end of the pterygoid 


196 Annals of the South African Museum. 


ramus, which has split into two thin horizontal plates—a long ventral 
one and a shorter dorsal one. Immediately behind the end of the 
basipterygoid process a Vidian foramen pierces the bone. It is on 
the latero-dorsal edge of the ventral pterygoidal plate that the epi- 
pterygoid stands; from this lateral edge a thin horizontal flange 
extends medially, and this acts as a base to the epipterygoid. From 
its base the epipterygoid stretches dorsally as a thin pillar, which 
is shown in broken lines in the figure. 

The pro-otic is a large bone; posteriorly, it is applied to the 
paroccipital process in the usual manner and forms the antero- 
dorsal part of the border of the fenestra ovalis; anteriorly, it stretches 
to the plane of the epipterygoid, which dorsally overlies its extreme 
antero-dorsal corner. The shape of the pro-otic is best understood 
from the figure; it forms nearly the whole of the lateral wall of the 
posterior part of the brain-case. A small foramen for the seventh 
nerve lies a short distance antero-dorsally to the fenestra ovalis; 
below it is a shallow depression for the geniculate ganglion; still 
further antero-dorsally the bone is pierced by a small rounded foramen 
for the sixth nerve, and under it is a depression; this foramen lies 
near the edge of a large opening through which the fifth nerve emerged; 
this opening appears to be a large oval foramen, whose anterior border 
is formed by the long anterior process of the pro-otic; it is not certain 
that this opening is bounded antero-dorsally by bone, as a part of 
the epipterygoid overlies this region; its general shape indicates a 
large oval foramen completely bounded by the pro-otic; the postero- 
dorsal border of this foramen is formed by a long process of the pro- 
otic, similar to the anterior one, which separates it from an equally 
large foramen lying dorsally to it; this is the large venous foramen. 

In anterior view, the edges of both pro-otics have been exposed 
and it is seen that, above the pituitary fossa, processes are sent 
inwards by both pro-otics, so as to form a bridge of bone over the 
pituitary fossa; ventrally, the bridge appears to be supported by a 
spur of the basisphenoid, which here forms the posterior border of 
the pituitary fossa; dorsally to this bridge, the pro-otics approach 
each other closely (3 mm.), so that the brain here emerged through 
a high and narrow slit. On the anterior face of the pro-otic, just 
above the bridge, is a small foramen facing directly forwards—this 
is for the exit of the fourth nerve. 

The sphenethmoidal region is unfortunately not preserved. 

The basioccipital has a well-developed condyle and has strong 
downwardly directed processes, on whose ventral surfaces the 


South African Gorgonopsia preserved in the British Museum. 197 


basi-sphenoidal tubera are applied; it forms the postero-ventral border 
of the fenestra ovalis; on its ventral surface lie the two nutritive 
foramina in their usual position. The foramen for the tenth nerve 
is overhung by the exoccipitals in the usual manner. 

If we are right in our interpretation that in Arctops the anterior 
part of the pro-otic is not ossified, and that the foramen for the fifth 
nerve is only represented as a shallow notch, and that in Cynariops 
the notch is deeper, but still unenclosed, then “‘Lycaenodon’”’ is a 
more highly developed form, as here the pro-otic has extended forward 
to surround the foramen for the fifth nerve. The type of Lycaenodon 
was shown to appear primitive, and it would thus seem that there is 
an additional reason for regarding this brain-case as not belonging 
to the genus Lycaenodon. The strong basioccipital, much stronger 
than in Cynariops, is, however, a primitive character, which may 
well be found associated with the characters found in the type of 
Lycaenodon. Until we know more about the relative values of 
characters styled “primitive” and “advanced,” any further expression 
of opinion will serve no useful purpose. 

B.M.N.H., R5746, Biesjespoort Station, Cape Province. 


Cynariops robustus, Broom. 
(Figs. 11-14.) 


The type consists of a good skull, lacking the temporal arches, 
part of the occiput, and the premaxillaries. After some further 
preparation, the lateral and dorsal surfaces show the sutures well; 
the relations of the various elements are best understood by referring 
to the figures. | 

The dental formula isi.5,c. 1 ,m.4; on the right side there are three, 
on the left four, the fourth being small; there is a diastema between 
the last incisor and the canine, whereas the first molar follows 
immediately on the canine. 

Mr. L. E. Parsons, Technical Assistant in the Geological Depart- 
ment, has prepared the palate and the lateral surface of the brain- 
case for me, and after some further development and etching, a 
number of interesting features have become apparent. The structure 
of the palate is adequately exposed; ventrally, the prevomers have 
a strong median keel and two lateral flanges on their anterior part; 
the median palatal groove has deep overhanging ridges, and both the 
palatine and pterygoid portions bear teeth; the median convoluted 
interpalatine suture is beautifully shown, and so is the transverse 


198 Annals of the South African Museum. 


suture separating the palatine and pterygoid in ventral view; the 
latter crosses the ridge at its lowest part and then continues in 
postero-lateral direction to meet the ectopterygoid suture. The 
posterior limits of the prevomers are not clearly shown, but probably 
are as indicated in the figure. 


Fic. 11.—Cynariops robustus. Type, B.M.N.H., R5743. Dorsal view of 
the skull. x 3. 


On the left side, the outer surface of the brain-case has been 
partially exposed; on this side, the epipterygoid is not preserved, 
but, on the right side, it is practically complete. The epipterygoid 
has a long base resting on the quadrate ramus of the pterygoid; it 
then extends dorsally as a very high, narrow, and flattened pillar to 
meet the parietal; it thus forms the side wall of the cavum epiptericum 
anterior to the pro-otic, with the Gasserian ganglion lying immedi- 
ately median to it; in lateral view, it lies in front of the lateral opening 
into the pituitary fossa and, further dorsally, overlies the anterior 


South African Gorgonopsia preserved in the British Museum. 199 | 


border of the large irregular notch for the fifth cranial nerve. The 
pro-otic is a fairly large bone, whose shape will be better understood 
from the accompanying figure than from a verbal description. 
Ventrally, it meets the basisphenoid in a fairly open suture; antero- 


La ie mice S 


B.Oc., B.Sph. 


Fic. 13.—Cynariops robustus. Type. Ventral view of the skull.. x &. 


ventrally, it forms the posterior border of the fenestra leading into 
the pituitary fossa; dorsally, it forms the ventral and posterior borders 
of the notch for the fifth nerve; the anterior pro-otic process is ap- 
parently not strongly developed, so that the notch is not deep; 
further dorsally, the pro-otic forms the ventral border of the large 


200 Annals of the South African Museum. 


P Ovb. Melee wae Deis 


Fig. 12.—Cynariops robustus. Type. Lateral view of the skull. x 4. 


Fie. 14.—Cynariops robustus. Type. Lateral view of the left side of the 
brain-case. x 4. 


South African Gorgonopsia preserved in the British Museum. 201 


venous foramen; but, unfortunately, the extent to which the supra- 
occipital and interparietal enter the side wall of the brain-case cannot 
be determined. The fenestra ovalis is seen to lie above and behind 
the basisphenoidal tubera, and appears to be fairly large. The 
foramen magnum is extraordinarily large and the basioccipital con- 
dyle is remarkably small. Unfortunately I have not been able to 
determine anything as to the presence of a parasphenoidal rostrum 
or the nature of the sphenethmoidal portion of the brain-case. The 
most striking feature about the brain-case of Cynariops is its great 
height and relative narrowness. In this respect it agrees with the 
fragmentary brain-case of “ Lycaenodon.”’ 

The main morphological features are: 

Primitive. Sloping occiput, not cupped; epipterygoid slender; 
preorbital depression; pro-otic of no great anterior extent. 

Advanced. Snout as broad as high and rounded; no step in 
maxillary border; orbits slightly anteriorly directed; pterygoid 
flanges anteriorly situated; temporal openings somewhat dorsally 
directed, but not large; parietal apparently enters brain-case; basi- 
occipital extraordinarily undeveloped; pro-otic larger than in 
Arctops; frontal forms only small part of orbital border; deep 
maxilla; short tooth row, but straight; parietal width reduced; 
paroccipital not massive. 

From this summary of characters, it is clear that Cynariops is much 
further advanced than Arctops, Gorgonops, Scymnognathus, and 
Leptotrachelus. In the nature of its brain-case it appears to occupy 
a position intermediate to Arctops and that represented by the 
brain-case referred to “‘Lycaenodon,” and it is in this respect com- 
parable to Scymnognathus and Leptotrachelus. These two latter 
genera are, however, in all other respects definitely more primitive. 

Chief measurements : 


From premaxilla to basisphenoidal tubera = 1A @ sani. 
From premaxilla to pineal foramen QUAL: 
From pineal foramen to edge of occipital plate . Ss 
From premaxilla to front of orbit . : é Gores. 
Interorbital width BOM 5: 
Intertemporal width . ; 2215; 
Width of snout . : Scans 
Height of snout . ; : Ae. 
Width across nierresidal anges BOP x 
Height of occiput . ; : ; : 45 ,, 


VOU. <kxI, PART 2. 1) 


202 Annals of the South African Museum. 


Length of molar series . : : ; : 19 mm. 
Diastema . : : : P : / De be 
Length of incisor series . ! 20 nae 


B.M.N.H., R5743, Biesjespoort Snort Outre Baume 


Lycosaurus pardalis, Owen. 


This form has been studied by Owen, Lydekker, Broom, and 
Watson. In Watson’s 1921 account all the evidence is fully stated. 
The left side is best preserved, and Watson’s figure is obviously a 
mirror image of this side. It is the left, and not the right, ramus of 
the mandible that is preserved. Although the difference in level 
between the pre-canine and post-canine edge is considerable, the step 
is not so abrupt as in Gorgonops. The dental formula is apparently 
1.5, ¢.1, m. 4%; the incisors are serrated, but as the teeth are badly 
preserved this is not well shown. The coronoid process of the dentary 
stretched far in dorso-posterior direction. 

The main morphological features are: 

Primitive. Orbits laterally directed; skull higher than wide; 
laterally situated nostril; slender epipterygoid. 

Advanced. Snout short, rounded but high; section of snout not 
square; no abrupt step in dentigerous border; maxilla deep; small 
facial part of septomaxilla. 

Although Lycosaurus is very inadequately known, it seems certain 
that it represents a stage somewhat in advance of that reached by 
the forms described above. As nothing is known of the basicranial 
axis and the posterior part of the skull, it would be best not to include 
this form in any morphological series. 

Chief measurements: 


From premaxilla to pineal foramen . 1202 mame 
From premaxilla to front of orbit . : 80e 
Length of lowerjaw . : . “sot 
Interorbital width : : SOM. 
Intertemporal width . : : : 2D ee 
Width of snout. : , : AS 
Height of snout . 2. ei a ae 
Height of mentum : 44 ,, 
Length of molar series . ; viele eae 
Diastema . : ; Guess 
Length of incisor series . ; Leh ase 


Type, B.M.N.H., R1717, Karroo, ee Province. 
iia, 0 or) ” 


ee a ib nama ne gts in 


South African Gorgonopsia preserved in the British Museum. 203 


A number of specimens have been referred to Lycosaurus, viz.:— 


49407. A badly weathered snout, which shows the dental formula 
1.5, ¢.1,m. %. Letjesbosch, Cape Province. 


49410. A weathered fragment of snout, which shows the dental 
Koumelarivo, €. I, me. dal 2. 


R512. Weathered fragment of snout. 


Arctognathus curvimola, Owen. 
(Figs. 15-17.) 


The type is a distorted and weathered skull; the occiput, roof of 
skull, and temporal arches are very much weathered; part of the 
palate had been developed, and the teeth of the upper jaw are well 
preserved on the right side. This type has been studied by Owen, 
Lydekker, Seeley, Broom, and Watson. Watson’s 1921 account is a 
detailed one, and the sutures of the dorsal and palatal surfaces are 
indicated in his figures. I have refigured the dorsal surface in order 
to show the relations of the septomaxilla and its foramen, the maxillo- 
nasal and the jugo-lacrimal sutures. The septomaxilla is a fairly 
large bone and the foramen is a fair-sized narrow slit—both larger 
than described by Watson. The dental formula isi. 4, c.1, m.5; the 
posterior edges of the canine and molars are serrated. 

The middle part of the palate was misinterpreted by Watson, who 
described the pterygoid as a large bone forming the roof of the median 
groove, carrying the two dentigerous ridges and reaching the pre- 
vomers and internal nares. The line of matrix lateral to the denti- 
gerous ridges, taken by Watson to be the pterygo-palatine suture, 
is a crack. The real suture is plainly visible as a thin, digitating line 
of matrix, which commences at the antero-median corner of the epi- 
pterygoid and then runs medially to cross the dentigerous ridge at 
its lowest part, then runs obliquely forward and continues in the 
median line as a well-marked wavy line of matrix. Posterior to the 
median groove, which has an abrupt hinder edge, there lies a distinct 
small interpterygoid foramen. 

The left side of the skull lacks the temporal arch, and I have 
succeeded in exposing the lateral surface of the brain-case from this side 
by removing the displaced portion of the lateral surface. This could 
have been done sixty years ago as it was a straightforward piece of 
development, which was accomplished by the use of a small chisel and 


204 Annals of the South African Museum. 


hammer only. As now developed, the skull shows all but the anterior 
quarter of the internal structure in parasagittal view. 


Ect. Peer. 


> Pter. 


—— 
— =~. 


Type, B.M.N.H., 47339. Ventral view 


of the skull, modified after Watson. x 2. 
Qu. = quadrate. Qu.Ju. = quadratojugal. 


Fic. 15.—Arctognathus curvimola. 


St. =stapes. 


The basioccipital condyle and the greater part of the paroccipital 
have been removed by a fracture; their nature is probably as indicated 


in broken lines in the figure. 


| 


South African Gorgonopsia preserved in the British Museum. 205 


The basisphenoid is present im toto. Ventro-posteriorly lie its 
tubera, which are thin and edge-like; they underlie the ventrally 
directed processes of the basioccipital and diverge in posterior direc- 
tion, so that the floor of the hinder end of the brain-case (for the 
cerebellum) is wide, and the fenestrae ovales far apart. Anteriorly, 
the basisphenoid extends as a thin vertical sheet of bone, which 
laterally carries the horizontal basipterygoid processes; the ventral 
part of this sheet forms the posterior part of the keel, which must 
further anteriorly be formed by the parasphenoid flanked by the 
pterygoids, but the exact relations of these elements cannot be 
determined as the fusion appears to be very intimate; the dorsal part of 
the basisphenoid is anteriorly confluent with the parasphenoid; its free 
anterior edge lies in the plane of the posterior border of the pituitary 
fossa and, in life, this edge must have supported the infundibulum; 
dorsally, the basisphenoid meets the pro-otic along a line running from 
the fenestra ovalis to the open side of the pituitary fossa. 

The outer surface of the pro-otic does not show up too well; 
posteriorly, it was apparently applied to the anterior face of the 
paroccipital in the usual manner; ventrally, it rests on the sides 
of the basisphenoid; anteriorly, its extent is not great; this is rather 
remarkable for a form otherwise showing many advanced characters; 
the anterior process of the pro-otic does not extend far, so that the 
exit for the fifth nerve appears to be only a shallow notch, whereas 
in “‘Lycaenodon”’ it is a large oval foramen; the outer surface of the 
anterior pro-otic process is deeply hollowed out, and this may have 
housed the Gasserian ganglion; dorsal to the opening for the fifth 
nerve, the hinder pro-otic process is also short, so that the venous 
notch has no extensive osseous ventral border. Dorso-anteriorly to 
the large fenestra ovalis lies a small foramen for the facial nerve, with 
a slight hollow under it for the geniculate ganglion. Anteriorly, the 
pro-otic forms the posterior border of the open side of the pituitary 
fossa and, posteriorly, forms part of the anterior border of the large 
and irregular fenestra ovalis. 

Dorsally, there are indications that both the parietal and inter- 
parietal sent down flanges, which contributed to the formation of the 
side-wall; the anterior part of this parietal flange articulated with 
the posterior part of the sphenethmoid; it was this relation that 
apparently led Watson astray, since, what he interpreted as a broad 
epipterygoid on the right side is, in reality, the outer surface of the 
sphenethmoid; the posterior part of the parietal flange articulated 
with the ascending epipterygoid. 


206 Annals of the South African Museum. 


The epipterygoid is not preserved, but there is no reason to suppose 
it to be different from that of the other known Gorgonopsia; standing 
on the pterygoid, it would ascend as a thin bony rod in a plane lateral 
to the other bones of the brain-case, and acted as a lateral wall for the 
thalamencephalon, mid-brain, and Gasserian ganglion. 


I 
ft 
sitar 


Fic. 16.—Arctognathus curvimola. Type. Dorsal view of the skull. x 4. 


The parasphenoid, from where it is indistinguishably fused with 
the basisphenoid, stretches antero-dorsally as a vertical sheet of bone; 
at its dorsal edge it appears to split, and in this groove the ventral 
edge of the sphenethmoid rests; anteriorly, the parasphenoid tapers, 
and here appears as a vertical slip of bone applied to the lateral surface 
of the sphenethmoidal part of the interorbital septum. 

The sphenethmoidal part of the brain-case is beautifully shown, and 
it is now evident that the “sphenethmoid ”’ figured in Scymnognathus 


ee 


South African Gorgonopsia preserved in the British Museum. 207 


and Leptotrachelus is only a part of that bone. The sphenethmoid in 
Arctognathus is a very large element; its ventral half represents a part 
of an ossified interorbital septum and rests on the parasphenoid, and, 
further forward, on the dorso-median keel of the pterygoid. Anteriorly, 
the interorbital septum is continued forward, but is separated by an 
unossified portion from the internasal septum. It is thus only the 
dorsal portion, which has split into two curved sheets of bone, that is 
preserved in Scymnognathus and Leptotrachelus. This housed the 


COP 


Fie. 17.—Arctognathus curvimola. Type. Lateral view of the left side of 
the brain-case. x 2. 


fore-brain. Dorsally, flanges from the frontal and parietal are sent 
downwards and these, meeting the dorsal edges of the sphenethmoid, 
complete the side-wall. Ventro-posteriorly, a large rounded foramen 
pierces the sphenethmoid; it is directed somewhat forward and out- 
ward, and from it may have emerged the ophthalmic branch of the 
fifth nerve, but the large size makes it more probable that it is really a 
venous foramen. Ventro-anteriorly, lies a downwardly-directed oval 
slit between the median septum and the side of the sphenethmoid, 
through which the second nerve emerged. Anteriorly, the sphe- 
nethmoid has a free edge, and anterior to this edge, the median septum 


208 Annals of the South African Museum. 


is visible. The olfactory lobes were probably situated in this position. 
Antero-ventrally to the tapering end of the parasphenoid, the sphe- 
nethmoidal part of the interorbital septum is extraordinarily thickened 
and, in this part, rests on the dorso-median keel of the anterior ptery- 
goid ramus. Thethickening is solid, and I can make no suggestion 
as to its probable function. Posterior to this structure there is a 
gap, which thus lies between the sphenethmoid, pterygoid, and 
parasphenoid. 

The general plan of all these internal ossifications in the skull of 
Arctognathus is very similar to that which is known in the Dicynodonts 
from Pearson’s account. 

The main morphological features may be summarised :— 

Primitive. Preorbital depression, but very weak; septomaxilla 
with large facial exposure and large foramen; epipterygoid apparently 
rod-like; fenestrae ovales far apart; pro-otic relatively small; 
fenestra ovalis not situated high up in the brain-case. 

Advanced. Snout short and rounded in section; nostril large; 
orbits not wholly laterally directed; small postfrontal; preparietal 
small or absent; short, deep maxilla; tooth row short; dentigerous 
border with a distinctive, ventrally convex curve, with no step; 
pterygoid flange anteriorly situated; interorbital width reduced; 
intertemporal width reduced, and fairly large temporal openings; 
basisphenoidal tubera reduced. 

Arctognathus is thus clearly a Gorgonopsian well advanced in the 
direction of development taken by this group, but it still possesses 
a number of primitive features. It has been shown that a number 
of features noted by Watson as advanced were cited on invalid 
grounds, viz. the septomaxilla and its foramen are large as in other 
Gorgonopsians and there is no evidence that the epipterygoid is 
broad. The pro-otic is certainly not very advanced, and stands on 
the same developmental niveau as do Scymnognathus and Lepto- 
trachelus; in this respect “‘Lycaenodon”’ is much more advanced. 
The nature of the basisphenoid and its relations with the pterygoids 
and parasphenoid do not show very marked advances over the 
condition of the earlier Gorgonopsians that have been described 
above. Inthe attainment of a short rounded snout and the reduc- 
tion of the interorbital and intertemporal width, however, Arctog- 
nathus shows definite advances. The great amount of ossification 
in the median septum appears to be primitive and will probably 
be found in the earlier forms when better preserved material is 
examined. 


South African Gorgonopsia preserved in the British Museum. 209 


Chief measurements: 


From premaxilla to basisphenoidal tubera ks arena. 
From premaxilla to pineal foramen my Ota, 
From pineal foramen to edge of occiput . ayy ps 
From premaxilla to front of orbit . : : gil). 
Length of lower jaw : LSS) 
Width across squamosals : ' ; 5 WAGY 
Interorbital width ; ; OMe 
Intertemporal width . ee En OOCn 
Width of snout . : G4. 
Width across lateral pervect anges ; : TA 96 
Height of snout . : A SON ®. 
Height of occiput DOONEY es 
Mentum : ; OY ee 
Length of molar series . ; : Sil 
Diastema . He 5 
Length of incisor series . TAB) ig 


Type, B.M.N.H., 47339, Kagaberg, Bedford, Cape Province. 


Arctognathus *curvimola, Owen. 


In the collection there is a preorbital portion of a skull with part of 
the lower jaw in position, but with the teeth badly preserved. Owen 
described it under the name Lycosaurus tigrinus; Broom, in 1911, 
showed that it could not be included in the genus Lycosaurus, and 
proposed the new name Arctosuchus, and, in 1932, considered it to be 
a Therocephalian. The specimen is a very bad type, but there is no 
doubt that it represents a Gorgonopsian, probably very close to 
Arctognathus. In 1921 Watson maintained that it was specifically 
identical with Arctognathus curvimola. I have etched the right side 
of the snout, and the maxillo-premaxillary suture now shows clearly; 
the relations of maxilla, premaxilla, septomaxilla, and septomaxillary 
foramen and the nostrils are typically Gorgonopsian and do not differ 
much from the condition in A. curvimola. There appear to be four 
incisors, although the roots of only two are actually preserved; the 
molars are badly preserved, but probably number five. All the 
features shown by the fragment thus agree very well with 4. curvimola 
and, as no differentiating features can be determined, there is no valid 
reason to distinguish it by name from A. curvimola. 

B.M.N.H., R1719, Mildenhalls, Fort Beaufort, Cape Province. 


210 Annals of the South African Museum. 


Cynodraco serridens, Owen. 


Owen described a number of fragmentary snouts under this name, 
and another fragment presumably associated with a good humerus 
under the name of C. major. There is no evidence to warrant the 
separation into two distinct species. Moreover, so little can be 
determined of the structure that, except for the deep mentum in one 
specimen, there is very little evidence that the fragments are Gor- 
gonopsians at all. Owen’s lithographs show the general appearance 
of these specimens very well. They are: 


Type, B.M.N.H., 47084, Bovey’s Farm, Fort Beaufort, Cape 
Province. 

This is a bad piece of snout, which exhibits some features of the 
incisors; there are five upper incisors with fairly strong and coarse 
serrations, four lower incisors with similar serrations; the incisors 
are strong and protruding. 


B.M.N.H., 47086, Fort Beaufort, Cape Province. 
A mandibulary fragment showing the canine of both sides and also 
four incisors. 


B.M.N.H., 47085, Stylkrantz, Graaff-Reinet, Cape Province. 

Weathered and fragmentary snout; the fragmentary incisors show 
fairly coarse serrations on both anterior and posterior edges. The spur of 
the septomaxilla protruding into the nostril is shown on the right side. 


B.M.N.H., 47309, Mildenhalls, Fort Beaufort, Cape Province. 

This is Owen’s type of.C. major. It consists of a fragment of 
mandibulary symphysis with both upper canines attached. The 
roots of four large incisors are seen in section; the serrations on the 
posterior border of the very large canines are beautifully preserved. 
The symphysis is deep. 

B.M.N.H., 47310, Mildenhalls, Fort Beaufort, Cape Province. 

A very good left humerus showing both foramina, well illustrated 
in Owen’s lithograph. 

Chief measurements : 


Length , : > 260 mime 
Width across the epicondyles : A - Dahon 

Width across the proximal surface . 105 by 36 mm. 
Width across delto-pectoral crest . - \LhOjmam: 
Maximum width of shaft . : 4 iranSoune 
Minimum width of shaft ’ IMRU2G 


Length of delto-pectoral crest : L of DASE ES, 


South African Gorgonopsia preserved in the British Museum. 211 


Cerdognathus greyi, Broom. 


The type is an imperfect dentary; the number of incisors cannot 
be ascertained, but four seems to be the most probable number; 
the canine and four molars are badly preserved as casts and, although 
no serrations can be seen, this evidence is simply negative; Broom 
is probably right in postulating a low coronoid process, as this would 
conform with the general contour. The very low, though upright, 
symphysis and the absence of a diastema stamps the dentary as a 
distinct type; but one wonders if such a fragment warrants the 
creation of a new generic name. 

Chief measurements : 


Probable length of dentary . ; a LOO? mm 
Height of symphysis. : : AM) og 
Length of molar series . : Si 4. 
Depth of dentary behind last molar ; i ae 


Type, B.M.N.H., R2892, Klippoort, Cradock, Cape Province. 


Scylacops capensis, Broom. 


This specimen consists of the middle third of a skull, which shows 
the greater part of the structure of that region. Watson’s description 
and figures appear to be correct in every detail. His attitude with 
regard to the specimen is to be highly commended. The snout is 
missing and the dentition is thus unknown and, as much of the classi- 
fication of the Gorgonopsia rests on this character, Watson, although 
recognising the fact that he had before him a representative of a 
new genus, refrained from naming it. Instead, he gave a good 
morphological account, which is of far greater value than a new 
generic name. Subsequent finds have shown the wisdom of such a 
procedure. We now have an excellent skull as the type of Scylacops 
capensis, named by Broom, but to Watson must go the honour of the 
first morphological description. 

B.M.N.H., 47098, ?Fort Beaufort, Cape Province. 


A Fore-foot of an Unnamed Form. 
(Fig. 18.) 


Lying on a series of vertebrae, there is a partial right fore-foot of 
a Gorgonopsian. The fourth digit is fully preserved; the third lacks 


212 Annals of the South African Museum. 


only the point of the claw; the fifth has two phalanges preserved; 
parts of the third, fourth, and fifth metacarpals are present; the 
fourth and fifth distals are present as a single fused element, to which 
the distal half of the ulnare is articulated. The foot is typically 
Gorgonopsian (about two-thirds the size of that of Aelurognathus 
tagriceps), with the distinctive mammal-like epiphysial distal ends 
to the metacarpals, and with indications of an incipient reduction 


Fig. 18.—An unnamed Gorgonopsian. B.M.N.H., R3768. Dorsal aspect 
of the partial manus. x #$. 


U. =ulnare. 
4+5 =fused fourth and fifth distal carpals. 
III-V =the third, fourth, and fifth digits. 


in the number of phalanges. The fourth and fifth metacarpals 
articulate with the fused fourth and fifth distals; the second phalanx 
of the fourth, and, in particular, of the third digit, is very much 
shortened; the third phalanx is robustly developed. 

As preserved, the phalangeal formula is—?, ?, 4,4, 3. If the first 
and second digits possessed two and three phalanges respectively, the 
structure of the digits would be exactly as in Aelurognathus tigriceps 
and, as in that form, dorso-ventral movement of the segments inter se 
is well developed with finely modelled articulatory faces. 

B.M.N.H., R3768, Oude Klip, Cape Province. 


South African Gorgonopsia preserved in the British Museum. 213 


Short Discussion. 


In a series of papers on the Therocephalians, Gorgonopsians, 
Bauriamorphs, and Cynodonts, Watson has argued that one branch 
of the Therocephalians gave rise to the Bauriamorphs, and that the 
Gorgonopsians led on to the Cynodonts. With the first conclusion 
I can concur, but in this study of a limited number of Gorgonopsian 
skulls two important facts bearing on this question have been 
established, viz. the nature of the epipterygoid and the relations of 
the posterior end of the prevomers, and these seem to invalidate 
Watson’s second conclusion. In all the Gorgonopsians I have 
studied there is no evidence, whatsoever, of a tendency for the original 
slender rod-like epipterygoid to become widened and intimately 
incorporated into the side-wall of the brain-case; and in all, where 
the palate is known, the prevomers have their posterior ends tapering 
and are here clasped by the palatines and do not meet the pterygoids 
on the ventral surface. In the Therocephalia, on the other hand, 
there is very definite evidence of the progressive widening of the 
epipterygoid and, in this group, the posterior end of the prevomers 
is expanded and underlies the pterygoid in ventral view. This is 
also the condition in the Cynodonts. With respect to the nature 
of the prevomers and the epipterygoid, it seems that the Gorgon- 
opsians must be excluded from the direct ancestry of the Cynodonts; 
but at this stage it would be premature to maintain that these two 
characters indicate a Therocephalian ancestry. I am more inclined 
to think that no known Therocephalian indicates anything more 
than a parallel development in the two suborders, Therocephalia and 
Cynodontia. I do not intend to continue this discussion any further; 
this can be more profitably done when the various suborders of the 
Theriodontia are treated monographically, as I hope to be able to do 
in the near future. 

My thanks are due to the officers of the British Museum (Natural 
History) for permission and facilities to study the material in their 
charge. I am particularly grateful to Dr. W. E. Swinton for his 
constant readiness to facilitate my work, and to Mr. Parsons for 
doing some of the developmental work. Professor D. M. 8. Watson’s 
interest was very encouraging, and his critical advice always welcome. 
To the University of Stellenbosch I am indebted for a small scholar- 
ship, which has enabled me to undertake the visit to London. All 
but three of the figures are by my wife. 


eo ig Ag ( 


ee ey deit J ai =e, «re: 
. ta. oo “i 
a x 9 ire Se ea hy 
= < ae Ae i> 
ae el ay rt 
3 ied ie ee 


Sta pot aan se caw 

; ry ee ee Pei ie adaghage wide 
wy E Parr ty & bea prt wea 
‘vt ited Sie : 

ed AD ate te aeeem ie 
bins hast atl = Ade meee 

, ae i -_ neem 


( 215 ) 


7. A Contribution to the Morphology of the Mammal-like Reptiles of 
the Suborder Therocephalia.—By Lizuwr D. Boonstra, D.Sc., 
Palaeontologist to the South African Museum and Queen Victoria 
Scholar of the University of Stellenbosch. 


(With 35 Text-figures.) 


Although many genera and species of Therocephalians have been 
described by Owen, Seeley, Broom, Watson, and Haughton since these 
forms became known about seventy years ago, very little is known of 
the cranial morphology. The outer surface of the skull has hitherto 
been the only structure adequately known; certain features of the 
palate of Scylacosaurus, Ictidosuchoides, Moschorhinus, Notosollasia, 
and Whaitsia have been described by Broom; Haughton has described 
the palate in Akidnognathus and Whaitsia, and a longitudinal section 
of the brain-case of Alopecognathus; Watson’s account of the palate 
and basicranial region of the Scaloposaurids has placed our knowledge 
of this family on a sound footing. 

In this paper the results of a study of the Therocephalians preserved 
in the British Museum (Natural History) are recorded. This collection 
consists of about twenty distinct types, and in a number of cases it has 
been possible to determine the structure of the palate and brain-case 
in addition to the dorsal, lateral, and occipital surfaces. As a number 
of these specimens are the historical types of Owen, Seeley, and 
Lydekker, a re-examination will serve as a useful introduction to a 
monographical account of the Therocephalia, which I hope to under- 
take in the near future. 

The suborder Therocephalia has hitherto been subdivided into five 
families, viz. Alopecopsidae, Ictidosuchidae, Pristerognathidae, Scalo- 
posauridae, and Whaitsidae; to these must now be added Huchamber- 
sidae and Lycideopsidae. 


The Pristerognathidae are represented in the collection by Alope- 
codon, Cynariognathus, Pristerognathus, Scylacosaurus, Scymnosaurus, 
and Trochosaurus; Hyorhynchus and probably also Theriodesmus may 
be included here. All these forms are from the Tapinocephalus zone, 
and represent the more generalised forms of the Therocephalia, which 
are on a definitely more primitive stage of development than the more 
advanced Scaloposaurids, Whaitsids, Huchambersia, and Lycideops. 


216 Annals of the South African Museum. 


Alopecodon cf. priscus, Broom. 


A very much weathered and badly preserved snout with the anterior 
third of the mandible, showing only the dentition in a very unsatis- 
factory manner, was, in 1925, made the type of a new species— 
Alopecodon minor, Broom. Broom gave the dental formula i. 7, c.3, m.7 
or 8. I find seven incisors as described by Broom; the two small 
teeth, which on account of their different direction may possibly be 
small canines, but may equally well represent two last incisors differing 
somewhat from the anterior ones, as there is no evidence that they are 
implanted on the maxilla; then a large canine, oval in cross-section; 
then a diastema of 8 mm.; then six closely packed molars followed 
by an empty socket, to which, medially, there lies a small tooth; the 
length of the molar series (7) is 24 mm. On none of the teeth are any 
serrations visible, but they may have been present in life. 

Both A. minor and A. rugosus are bad types. Broom’s specific 
characters comprise only the number of teeth, and as these are badly 
preserved, it is not a reliable criterion. In A. priscus the dental 
formula given by Broom isi. 7,¢.2+1,m.8?; in A. minori.7,c.2+1, 
m.7or8; in A.rugosusi.?,c. ?+1,m.7. Considering the nature of the 
preservation in these three specimens, the dental formulae approxi- 
mate so closely that, in the present state of our knowledge, it would 
appear advisable to consider both A. minor and A. rugosus as examples 
of Alopecodon priscus. It is true that they are considerably smaller 
forms, but this does not appear to be a sufficiently weighty reason for 
retaining three distinct specific names, each of which is only repre- 
sented by a very incomplete skull. 

B.M.N.H., R5750, Abraham’s Kraal, Prince Albert, Cape Province. 


Cynariognathus platyrhinus, Broom. 
(Fig. 1.) 

This specimen is a weathered snout, which Broom has determined 
as belonging to the same species as the type snout now in the 
American Museum. Sufficient of the teeth are preserved in a fair 
state of preservation to enable one to determine something of the 
dentition; two anterior and two posterior incisors and a space for 
another in between are present; there is one large canine, oval in 
cross-section; then a series of six closely set, fairly small molars; 
posterior to the sixth, there may have been one or more additional 
molars. In the lower jaw, a smaller canine followed by eight very 
closely set molars are visible; there is no diastema as in the maxillary 


Morphology of Mammal-like Reptiles of Suborder Therocephalia, 217 


series; there does not seem to be room for more than three lower 


1 2 
incisors. The dental formula would thus be - ee nS ere * For 
ie 2,0 deanma8 


the type, Broom has given the P 
1,6, Go ls in. & Laie 


i.3,c.1,m.8 BY 
GC 


following formula : 


As both specimens are not well 
preserved, I believe it legitimate 
to assume that in life the denti- 
tions were probably similar. In 
the British Museum specimen the 
posterior border of the incisors, B. 
canines, and molars all carry fine 
serrations. Of the type, Broom 
stated, “There is no evidence of 
any serrations, and were they 
present the specimen would be A. 
expected to show some of them.” 
If Broom’s observations are cor- 


rect, there are thus points of Fic. 1. — Cynariognathus platyrhinus. 
B.M.N.H., R4097. Diagrammatic 


difference between the two speci- cross-sections through the snout. 
mens. Further evidence, based x 1. 
on more complete and better pre- ~ At the level of a kas 

: ; : ss ne ourth ,, 
served specimens, is, however, C. iM  thind & 


necessary before a new species Mx.=maxilla. Pr.V. =prevomer. 
can legitimately be created. Pal.=palatine. Pter. =pterygoid. 

I have had three slices cut across the anterior part of the snout. 
These are reproduced here ina slightly diagrammatised form. It is seen 
that, posteriorly, the prevomers are fused, whereas, anteriorly, the 
fusion becomes less intimate and finally ceases; the palatine is seen 
to form a paired dorsal keel in the median line, which is supported by 
a girder of the anterior pterygoid ramus. Through the weathered 
right side the dorsal paired keel of the prevomers is seen; this 
supported the soft internasal septum. 

B.M.N.H., R4097, Uitkyk, Prince Albert, Cape Province. 


Pristerognathus polyodon, Seeley. 


The specimen in the collection is the type of the genus; it consists 
of a weathered snout, from which very little of morphological interest 
can be determined. As far as can be ascertained the dental formula 


WOllis LOO, INE AB 20 


218 Annals of the South African Museum. 


i, Ole oa ema 
3) Os lL ita OF 
fine serrations; in the upper incisors the crowns are lost, but one may 
assume that they also were serrated; the posterior border of the upper 
canine (and lower?) carry somewhat coarser serrations. The incisors 
are fairly large—so is the canine; the molar roots are of the same size 
as those of the incisors. There is a step on the maxillary border 
anterior to the canine. The splenial practically enters the symphysis 
on the ventral surface; further dorsally, it may actually enter it. 
Chief measurements: 


is : The posterior border of the lower incisors bears 


Length of the six upper incisors . : . 35 mm. 
Length of the three lower incisors ! -, | ae 
Width of upper canine ; is, See 
Width of the snout . : ; “+ 255i 
Height of the snout . ' ; Morr 


Type, B.M.N.H., R2581, Cypher, Tamboerfontein, Beaufort 
West, Cape Province. 

Under the number R5753 there is registered a badly preserved 
shoulder girdle, of which Broom has given good restored figures under 
the name ?Pristerognathus minor. The number R5752 refers to some 
cervical vertebrae of the same form. 


Scylacosaurus sclatert, Broom. 
(Figs. 2-5.) 

A nearly complete, though slightly weathered, skull, preserved in 
an extremely intractible matrix, was purchased by the British Museum, 
in 1912, from J. H. Whaits. This specimen has been referred to the 
above species. In point of size and arrangement of the external bones 
of the snout it agrees exactly with the type in the South African 
Museum. Broom’s account (1903) of the dentition of the type gave 
the dental formula 1. 6, c. 3, m. 7, incisors without serrations, canine 
probably with serrations. In 1932, he gave the formula 1. 6, c. 2, m. 7. 
In this specimen the incisor series is composed of five small, slender, 
and closely packed teeth; these are definitely on the premaxilla; 
behind them are three much smaller teeth; these appear to lie on the 
maxilla, as in lateral view the premaxillo-maxillary suture lies anterior 
to them; posterior to a short diastema lies the large canine 
(20x 7mm.); the molar teeth are of the same size and nature as the 
anterior incisors; on the right side, a closely set series of seven teeth 


sete 


Sere 


Fig. 2.—Scylacosaurus sclateri. 


xs 


B.M.N.H., R4055. Dorsal view of the 
skull. 


Abbreviations to this and the subsequent figures. 


Ang. =angular. 

Art. =articular. 

B.Oc. =basioccipital. 

B.P.P. =basipterygoid processes. 
B.Sph. =basisphenoid. 

Cor. =coronoid. 

Dent. =dentary. 


ict. Pter. =ectopterygoid. 
Ep.Pter. =epipterygoid. 


Ex.Oc. =exoccipital. 
Fen.Ov. =fenestra ovalis. 
BL. =foramen incisivum. 
FJ. =foramen jugulare. 
Ie. =frontal. 

I.Na. =internal nares. 
Ju. =jugal. 

La. =lacrymal. 

Mx. = maxilla. 

Na. =nasal, 

Pa. = parietal. 

Pal. = palatine. 

Jee dite = postfrontal. 

Pin. =pineal foramen. 


Pit. 
P.Oc. 
P.Orb. 


Pr. Art. 


Pr.Fr. 


Pr.Mx. 


Pr.Ot. 


Ven. 


= pituitary foramen. 
= paroccipital. 

= postorbital. 

= prearticular. 

= prefrontal. 

= premaxilla. 

= pro-otic. 

= prevomer. 

= parasphenoid. 

= pterygoid. 
=post-temporal fenestra. 
= quadrate. 

= quadratojugal. 
=surangular. 
=septomaxillary foramen. 
=septomaxilla. 
=supraoccipital. 
=suborbital vacuity. 
= splenial. 
=squamosal. 

= stapes. 

= tabular. 

= venous foramen. 


Roman numerals refer to the cranial nerves. 


220 Annals of the South African Museum. 


occupy 23 mm.; on the left side, there are six less crowded teeth, 
with a gap for one additional tooth, occupying 27 mm.; the dental 
formula thus appears to be 1.5, c.1+3,m.7. All the teeth bear fine 
serrations on the posterior border. The dentigerous border curves 
upwards anterior to the canine, but there is no step. There are thus 
some differences in the dentition between the type and this specimen; 
allowing for the state of preservation and for a certain amount of 
individual variation, these differences should not restrain us from con- 
sidering them specifically identical. 

Mr. HK. L. Parsons, at my request, attempted to display the palate 
and occiput, but found that the bone was more fragile than the hard 


S.Mx. Na. Pelr Fro PFEr POrh Ja. 


Fic. 3.—Scylacosaurus sclateri. B.M.N.H., R4055. Lateral view of the skull. x ?. 


matrix, and had to abandon the attempt. On taking up the specimen 
again, I decided to make another attempt, and have succeeded in 
sufficiently exposing the left side of the brain-case and parts of the 
posterior half of the palate, so as to determine the general structure. 
In the accompanying figure, the anterior part of the palate has been 
drawn from the type-specimen in the South African Museum, as 
figured by Broom, and the posterior half from the British Museum 
specimen. The chief points of interest are: the widening of the 
basisphenoid and the bulge on the paroccipital, which characters are, 
in some respects, paralleled in the Scaloposaurids; the prevomers 
have a posterior shovel-shaped expansion, which underlies the ptery- 
goid; the anterior palatal fenestrae are relatively short; the paired 
tooth-bearing ridge lies wholly on the pterygoid, not, as in the Gor- 


Morphology of Mammal-hke Reptiles of Suborder Therocephalia. 221 


gonopsians, partly on the palatine; the suborbital vacuities are large; 
an interpterygoid vacuity is present; the pro-otics enclose a broader 
space than in the Gorgonopsians; the auditory groove, formed by a 
ridge of the squamosal, is well developed. 

The brain-case does not show up very well, but is of considerable 
interest in that it shows that there is a certain amount of parallelism 
between this Pristerognathid 
and the Scaloposaurids, and Pr Me 
in that it differs considerably te 
from that of another Pristero- 
gnathid—Trochosaurus. 

The basisphenoid is a 
strong bone with widely sepa- 
rated, ridge-like  tubera; 
lateral to the tubera proper, 
the basisphenoid extends 
still further laterally, and pon ye 
here forms a sharp distinct- Pee © Nowe ct. Phor 
ive ridge, which bounds AB : 
the fenestra ovalis antero- 
laterally and then continues 
posteriorly to meet the par- 
occipital (figs. 4, 5). This 
sharp ridge is not known in 
any other Therocephalian, 
but is present in Galesaurus 
andsome Anomodonts. An- 
teriorly, the basisphenoid 
carries a ventral keel and, 

: Bike 
anterior to the plane of the Qu 
pituitary fossa, carries the Fic. 4.—Scylacosaurus sclateri. B.M.N.H., 
horizontal basipterygoid FOS. Ventral view ofthe hl 
processes on its lateral face. 
Nothing can be determined of the relations of the basisphenoid with 
the parasphenoid. 

The lateral development of the basisphenoid, mentioned above, 
also occurs in the Scaloposaurids and Bauriamorphs, but if Watson’s 
account is correct, the relations differ somewhat, e.g. the flange does 
not bear a sharp ridge and, in Watson’s figure of Scaloposaurus, the 
fenestra ovalis lies anterior to the paroccipital and lateral to the 
widened end of the basisphenoid, whereas in Scylacosaurus it lies in 


222 “Annals of the South African Museum. 


the normal Therapsid position between the basisphenoid, pro-otic, 
paroccipital, and basioccipital; in Scaloposaurus there does appear 
to be a hollow filled with matrix in the same position as the fenestra 
in Scylacosaurus, and, moreover, the depression figured by Watson 
as the fenestra ovalis in Scaloposaurus also occurs in Scylacosaurus, 
and here it definitely is not the fenestra ovalis. 

The pro-otic is strongly developed; its anterior margin is overlain 
by the epipterygoid so that the cavum epiptericum has not yet been 
included in the cranial cavity; the ventral part of this bone has been 
removed and it can now be seen that -the posterior border of the 
lateral opening into the pituitary fossa is formed by the basisphenoid 


Ply Prk. Fin. 
—_ 


| Nee 


= 


Fig. 5.—Scylacosaurus sclatert. B.M.N.H., R4055. Lateral view of the left side 
of the brain-case. Occiput, basipterygoid process, and the postorbital bar 
seen in section. Lower half of epipterygoid removed. x l. 


and pro-otic; in lateral view the epipterygoid obscures the perforation 
for the fifth cranial nerve; further dorsally les the large venous fossa 
found in all Therapsids; the dorsal limits of the pro-otic cannot be 
determined; here the side-wall is formed by flanges from the parietal, 
interparietal, and supraoccipital, but their exact relations cannot 
be ascertained. As far as can be determined, the anterior margins 
of the pro-otics do not approach each other as they do in some 
Gorgonopsians. 

The epipterygoid is a high bone standing with its elongated base 
on the quadrate ramus of the pterygoid in a plane immediately 
lateral to that occupied by the pro-otic, forming a cavum epiptericum; 
it is fairly narrow and flattened, but is proportionately twice as 
broad as in any known Gorgonopsian, and roughly half as broad as 


Morphology of Mammal-like Reptiles of Suborder Therocephalia. 223 


in the Whaitsids and Cynodonts; dorsally, its inner surface is applied 
to the parietal, and its outer covered by a flange of the postorbital. 
The lower jaw is of the usual Therapsid nature, and its structure 
is best understood by referring to the figure, where it is shown in 
lateral view. 
Chief measurements : 


Length from premaxilla to the basioccipital condyle 2) doen 
Length from premaxilla to the pineal foramen MLNS) 38) 
Length from pineal foramen to edge of occipital plate . 25 ,, 
Length from premaxilla to anterior margin of orbit ue noo: o 
Length of the lower jaw : ; ; oa HOO! 5 
Width across the squamosals 93. 5 
Interorbital width : ; : 221 ae 
Intertemporal width . ! : : os 
Width of the snout. , : : Sob ee 
Height of the snout. ‘ : ; ; : S00 
Height of the occiput . : ah ibe Oy as 
Width across the lateral pteryaoid ames OW Ais 
Height of mentum of lowerjaw . £4, EOD as, 
Length of the molar series . : . Zi. 5, 
Diastema . : ; d ; : " : Ss. 
Length of the incisor series . s : : ' 4 Die 


B.M.N.H., R4055, Fraserburg Road Station, Prince Albert, Cape 


Province. 


Scymnosaurus watsom, Broom. 
(Figs. 6-8.) 

The type-specimen consists of the greater part of a large skull, 
lacking the snout; some vertebrae and ends of limb-bones, from 
which very little can be determined. The palate was figured, in 
1914, by Watson under the name Lycosuchus?. In 1915 Broom 
published outline drawings of the dorsal and lateral aspects, without 
indicating the constituent bones, and proposed the new name— 
Scymnosaurus watsont. In 1921 Watson published a fuller account; 
his figures and account of the internal aspect of the brain-case I find 
to be correct, and here he corrected his former error in the orientation 
of the articulatory surfaces for the lower jaw. I have refigured the 
palatal aspect, as my interpretation of its structure differs con- 
siderably from that of Watson. The most obvious error is that he 
maintained that a median bone, which he called the vomer, was 


224 Annals of the South African Museum. 


present in the middle third of the palate, lying between the pterygoids 
and palatines, and posteriorly separated from the basisphenoidal 


Fia. 6.—Scymnosaurus watsoni. B.M.N.H., R4100. Dorsal view of the 
Skalle) = xt 


rostrum by the interpterygoid vacuity and the pterygoids. The 
long median slip of bone interpreted as ‘“‘vomer,” is really the fused 


Fia. 7.—Scymnosaurus watsoni. B.M.N.H., R4100. Lateral view of the 
skull. x 4. 


dorsal keel of the pterygoids, on which the median septum rested. 
The state of preservation of the specimen is, however, such that it 
is only in the light of our increased knowledge that we are able to 


Morphology of Mammal-hke Reptiles of Suborder Therocephalia. 225 


establish the fact that no median “vomer” can possibly be present 
in this position. From my figure it can be seen that the relation 
of the prevomers, palatines, pterygoids, and ectopterygoids is typical 
as in all known Therocephalia. 

Attention may here be drawn to the points in which the Thero- 
cephalian palate differs from that of the Gorgonopsians. 


ee ae 
= 


sic 


Tub 
Fie. 8.—Scymnosaurus watsoni. B.M.N.H., R4100. Ventral view of the 
skull. x 4. 


(a) The internal narial opening is relatively shorter and, in the 
Whaitsids, is wholly or partly bipartite. 

(6) Posterior to the plane of the nares, the prevomers expand in 
shovel-shaped fashion and underlie the palatines and ptery- 
goids, whereas in the Gorgonopsians the prevomers end 
posteriorly as tapering slips of bone, laterally flanked by 
the palatines. 

(c) The suborbital vacuities are large, and consequently the ecto- 
pterygoids are reduced to form their posterior girder-like 


border. 


220; Annals of the South African Museum. 


(d) The median dentigerous ridges, when present, are carried wholly 
by the pterygoids. 

(ec) The pterygoid has a long anterior ramus, which meets the 
prevomers; the palatines thus do not meet in the median 
line, whereas they do in the Gorgonopsians. 

(f) The quadrate ramus of the pterygoid has a straight outer edge, 
whereas in the Gorgonopsians it is concave. 

(g) The paroccipital bar is a much stronger element. 

(hk) The ventral basisphenoidal keel is very deep in nearly all 
Therocephalians. 


I had considerable difficulty in determining the structure of the 
dorsal and lateral surfaces, but, by a lengthy process of alternately 
grinding down with a file and etching with acid, the limits of the 
majority of elements are now visible, and are shown in the accom- 
panying figures. The main points of interest are: the deep preorbital 
depression; the two grooves on the frontal, which is short and 
cruciform with only a small entry into the orbital border; the large 
prefrontal and postfrontal; the deep maxilla; the peculiarly high 
slip of the jugal on the postorbital bar; the high, thin parietal crest; 
the somewhat anteriorly directed orbits; the small posttemporal 
fossae; the epipterygoid, with straight edges, proportionately twice 
as broad as in the Gorgonopsians. 

Chief measurements : 


Length from premaxilla to the basioccipital condyle . 240? mm. 
Length from premaxilla to the pineal foramen : . MSO Fare 
Length from pineal foramen to the occipital plate . « Oma 
Length from premaxilla to the front of the orbit . . Oe 
Length of the lower jaw : F , LOD aes 
Width across the squamosals ; : : : , 2a 
Interorbital width : ‘ : : : : 50m 
Intertemporal width . : : ‘ eae : homes 
Width of the snout. ; : : : Geb 5. 
Width across the lateral nterroeid Toe 5 SEINE 
Height of the snout. : ; 62a 
Height of the occiput : - LOD ea 
Length of the molar series : Doe 


Type, B.M.N.H., R4100, Uitkyk, Prince Albert, Cape I Province. 


Morphology of Mammal-like Reptiles of Suborder Therocephalia. 227 


Trochosaurus major, Broom. 
(Figs. 9-11.) 
The type of this species is in the American Museum, and was 


described by Broom as Trochosuchus major. Haughton described 
a specimen in the South African Museum as Trochosaurus intermedius, 


Wc 
Fie. 9.—Trochosaurus major. B.M.N.H., R5747. Ventral view of the 
skull. x 4. 


and, in 1932, Broom referred his species to Haughton’s genus. The 
specimen in the British Museum formed the subject of the figures of 
the dorsal and lateral surfaces published by Broom in 1932. 

After Mr. Parsons had done the preliminary development, I succeeded 
in exposing the whole palate, occiput, and the greater part of the 
lateral surface of the brain-case on the right side. 

In the type, Broom found the dental formula to be 1.5, c. 2, m. 3%; 


228 Annals of the South African Museum. 


the incisors have a strongly serrated posterior border. In the British 
Museum specimen the dental formula is identical, but here there is, 
in addition, definite evidence that the molars as well as the canines 
and incisors are replaced; it is not certain that the last incisor is 
implanted on the premaxilla. 

I find Broom’s figures of the dorsal and lateral surfaces correct; 
in this supplementary account only the palate, occiput, and lateral 
aspect of the brain-case will be figured and described. 

The skull had been fissured, and subsequently weathered at the 
plane of the lateral pterygoid flanges so that this region is not very 
well shown; the middle part of the palate is revealed by a frontal 
fracture; the anterior and posterior portions have been chiselled out. 

As shown in the figure, the palatal structure is typically Thero- 
cephalian; the paired prevomers separate two short and wide 
fenestrae—the anterior part of which housed the lower canine, and 
the posterior functioned as internal nares; the prevomers have a 
shovel-shaped posterior expansion, which underlies the palatines 
and pterygoids; the palatines do not meet in the median line; the 
suborbital vacuities are large, and the ectopterygoids appear to be 
beam-like structures forming the posterior and lateral borders of 
these vacuities; the extent to which the ectopterygoids participate 
in the formation of the transverse pterygoidal bar cannot be deter- 
mined; posteriorly, the deep median keel, formed by the pterygoids 
flanking the basisphenoidal rostrum, is a prominent feature; the 
quadrate ramus of the pterygoid has a nearly straight outer edge; 
the basisphenoidal tubera are very massive and rounded, and diverge 
greatly and have a deep hollow between them, with the result that 
the fenestrae ovales are very far apart and the stapes short; there 
appears to be no evidence of a “‘vomer”’ underlying the basisphenoidal 
tubera. The flat nature of the anterior part of the palatal roof is 
in strong contrast to the deeply vaulted antero-median part of the 
palate of the Whaitsids. 

In the occiput the structure is adequately shown, only the basi- 
occipital condyle being lost. In general appearance the low and 
wide occiput is very similar to that of the Whaitsids, but there are 
great differences in the proportions of the constituent elements. 
A curious step above the supraoccipital forces the interparietal on to 
the dorsal surface; the post-temporal fenestrae are situated very 
high up and are far apart; the paroccipitals are enormously developed, 
their distal ends being bipartite, the lower supporting the quadrate 
and the upper supporting the strongly developed auditory groove 


Morphology of Mammal-like Reptiles of Suborder Therocephalia. 229 


of the squamosal. In posterior view, the quadrate and quadratojugal 
appear to be unfused, the quadrate is rounded and forms the main 
articulation, whereas the quadratojugal is exposed on the lateral 
surface and forms the small outer part of the articulatory surface; 
the stapes is short; the foramen magnum is small; in section it is 
seen that, at the plane of the foramen magnum, the dorso-lateral 
corners of the condyle are formed by the exoccipitals; the foramen 
jugulare is large; the basioccipital sends two processes ventrally to 
support the widely separated and massive basisphenoidal tubera; 
on this specimen I can see no evidence that there are three bones in 
this region; if, however, these ventrally directed processes really 
represent the basisphenoid intimately fused to the basioccipital, then 
the bone lying ventrally to them would represent a parasphenoid. 


Fig. 10.—Trochosaurus major. B.M.N.H., R5747. Posterior view of the 
occiput. x %. 


A successfully induced parasagittal fracture has enabled me to get 
at the brain-case from the right side; this section also reveals the 
structure of the occiput (fig. 11). 

The basisphenoid is seen to be a large bone with very strong 
posteriorly directed tubera, which rest on the basioccipital processes; 
the floor of the posterior part of the brain-case is thus very wide; 
laterally, it forms the ventral and anterior borders of the fenestra 
ovalis (the stapes is in situ), and its rostrum is flanked by the ptery- 
goids; the basipterygoid processes are horizontal and flattened, and 
are immovably clasped by the pterygoids; dorsally, it meets the pro- 
otic in a long suture and forms the lateral border of the opening 
leading into the pituitary fossa; anteriorly, it is continued as a 
median keel, but it has not been possible to determine where the 
transition into the parasphenoid occurs; the pituitary would le in 
the dorsal surface of the basisphenoid. 


230 Annals of the South African Museum. 


The pro-otic is a large bone, whose anterior portion is flanked by 
the epipterygoid; posteriorly, it is applied to the anterior surface of 
the paroccipital, and in part to the squamosal; antero-dorsal to the 
fenestra ovalis lies a small foramen for the seventh cranial nerve, and 
under this there is a small depression for the geniculate ganglion; as, 
in lateral view, the anterior part of the pro-otic is overlain by the 
epipterygoid, forming a cavum epiptericum, the foramina for the fifth 
and sixth nerves cannot be seen; dorsally, the pro-otic forms the 
ventral border of the venous fossa. 

Dorsal to the pro-otic, the side-wall of the posterior part of the brain- 
case 1s formed by the supraoccipital, interparietal, and parietal; the 


! ROD) 


Ep. Pler. 


Fic. 11.—Trochosaurus major. B.M.N.H., R5747. Lateral view of the right side 
of the brain-case. Occiput and postorbital bar seen in section; part of the 
epipterygoid removed to show the pro-otic. x 4. 


relations of these bones are shown in the accompanying figure and 
requires no verbal description. 

The epipterygoid is a large and broad plate of bone, which forms the 
side-wall of the cavum epiptericum; ventrally, it stands on the 
quadrate ramus of the pterygoid in the usual Therapsid manner: 
continuing upwards it obscures the lateral opening into the pituitary 
fossa and the foramina for the fifth and sixth nerves from lateral 
view; dorsally, it meets the parietal and is flanked by a thin sheet of 
the postorbital; the epipterygoid is thus seen to be relatively more 
than twice as broad as that of any known Gorgonopsian, and has the 
appearance of the relations hitherto known only in the Cynodonts, 
amongst the Therapsids. It differs from that known in Scymnosaurus, 
Scylacosaurus, and Ictidosuchoides in that it is not a relatively narrow 
plate of bone with straight edges, but has a shaft-like middle and 


Morphology of Mammal-like Reptiles of Suborder Therocephalia. 231 


expanded upper and lower ends, a condition reaching its greatest 
development in the later Whaitsids. 

In the nature of the brain-case and the basicranial axis, T'rocho- 
saurus differs considerably from that of the other members of the 
family Pristerognathidae. An examination of further material will 
undoubtedly show that this family includes forms which will event- 
ually have to be arranged in more than one group. It is, moreover, 
clear that the Therocephalians are, already in the Tapinocephalus zone, 
a well-established group, in which a number of different evolutionary 
tendencies have been developed. 

Chief measurements: 


Length from the premaxilla to the basisphenoidaltubera . 225 mm. 


Length from the premaxilla to the pineal foramen . ot ROME S 
Length from the pineal foramen to the edge of the 

occipital plate : oy ae 
Length from the premaxilla tp étie Hone of He osbit 5 I 
Length of the lower jaw “ . , SheoiliGe us. 
Width across the squamosals 4 ; beh GOL Oe: 
Interorbital width : . 402 
Intertemporal width . ‘ : : : ; 5 20m 
Width of the snout. . : Bb, 
Width across the lateral pterygoid flange : : PRO O Le 
Height of the snout . oe GOR: 
Height of the occiput . cihwaieO) Bes 
Length of the molar series. ; Area bali 
Length of the diastema : ae 
Length of the incisor series . : inte! 0 a 


B.M.N.H., R5747, Tapinocephalus zone ?, South Africa. 


Hyorhynchus platyceps, Seeley. 

The incomplete skull shows the structure of the middle third of the 
roof of the skull; the frontal is short and has a large entry into the 
orbital border; the postfrontal and postorbital are well developed; 
the pineal foramen is large; the parietal region is narrow. Although 
some bones are visible on the palatal surface, I do not quite understand 
their relations. Hyorhynchus is undoubtedly a member of the Pristero- 
gnathidae, and the few characters that can be determined seem to 
indicate a close relationship to Pristerognathus. 


B.M.N.H., R872, Gouph, South Africa. 


232 Annals of the South African Museum. 


The Ictidosuchidae are represented in the collection by two speci- 
mens referred to two genera, viz. Cerdodon and Ictidosuchoides. 
Cerdodon, which is not well known, is from an uncertain horizon in 
the Lower Beaufort, whereas Ictidosuchoides is known from a good 
skull, and is from the Endothiodon zone. 


Cerdodon tenuidens, Broom. 

In 1878, T. Bain had collected a calcareous nodule, which contained 
part of a Therapsid skull; the anterior part of the left side was 
weathered .and showed a portion of maxilla, premaxilla, and dentary. 
So little could be determined that neither Owen, Seeley, Lydekker, or 
Watson named it, but, in 1915, Broom produced a generic description, 
together with a number of others equally valueless. It is a pity that 
some authors will persist in naming such fragments, which are of no 
morphological or phylogenetic value whatsoever. 


1.5.%,¢. 1, m. T=82 y 
In 1915, Broom gave the dental formula, ———_—___—____, and in 
1.5; €. lane 
62s ce 1 mae) 1.4, c. i 4 
1932, DOP ee . I find the formula to be, 1 Se The 
1. te ol mee 1. 35 (Cs. aes 


posterior border of the lower molars, which alone have the crowns 
preserved, are serrated; the incisors are small and slender. 

I asked Mr. Parsons to attempt to have a parasagittal section made 
in order that something more definite may be known of this “‘type.”’ 
Unfortunately, this section was not cut true, with the result that very 
little of the structure could be determined; an additional slice was cut 
in an attempt to obtain a section along the median line, but also 
without success. The external weathered surface of the left side, 
figured by Broom, is thus all that can be determined in this new genus. 

Type, B.M.N.H., 49420, Gouph, South Africa. 


Ictidosuchoides longiceps, Broom. 
(Figs. 12-14.) 

In his original description Broom gave the dental formula, 1. ?, c. 3, 
m. 9, and maintained that anterior to the large canine there were an 
additional two teeth implanted in the maxilla. Unable to find these 
two teeth, and thinking that they may have been lost in transit and 
through handling, I ground down the anterior maxillary border in 
order to expose the roots. At the level of the maxillary edge two 
filled alveoli were found, but no crowns extended beyond the maxillary 
border; it would thus appear that these two teeth were in the process 
of eruption. No serrations are visible on the canine or the molars. 


Morphology of Mammal-like Reptiles of Suborder Therocephalia. 233 


Broom found that due to the narrowness and depth of the anterior 
part of the palate the infilling matrix could not be removed without 
damaging the maxillary teeth. By careful preparation with a fine 


f=, - i-S 2 
‘ 7 ost i ee — ve 
0 A Eee ny " ‘ " 
1 ‘ Oo cee 7 “ 
soe = ; 
Ko at Ae “ , Ste Me oe 
Sac Jd =e Os 3 , 
ose s 
ee, 4 


Fie. 12. — Ictidosuchoides longiceps. Fie. 14. — Ictidosuchoides longiceps. 
B.M.N.H., R5744. Dorsal view of B.M.N.H., R5744. Ventral view 
the skull. x 4. of the skull. x 3. 


Fig. 13.—Ictidosuchoides longiceps. B.M.N.H., R5744. Lateral view of the 
skull. x 4. 


chisel I have been successful in exposing the whole palate, but, un- 

fortunately, the premaxillaries had fallen out prior to fossilisation, 

and thus the nature of the junction of the palatal processes of the pre- 

maxillaries with the prevomers cannot be determined. An oblique 

fracture at the level of the last molar reveals that, on the dorsal surface 

of the unfused prevomers, a pair of ridges are developed, and the groove 
WO, VOOR, DAI A, 21 


234 Annals of the South African Museum. 


in between them must have supported the soft internasal septum. The 
prevomers are of the usual Therocephalian type; anteriorly, the end 
is fan-shaped, and posterior to the level of the internal nares, they 
again expand and underlie the anterior rami of the pterygoids and the 
median edges of the palatines well behind the level of the last molar. 

The interpterygoid vacuity has its posterior margin somewhat 
indefinitely preserved, but it would appear that it was very similar to 
that of Scaloposaurus and Icticephalus. The suborbital vacuities are 
large, and the lateral border formed by the ectopterygoids are of the 
narrow beam-like character as in Scaloposaurus. The sweep of the 
quadrate ramus is typical of the Therocephalians, and nothing of the 
web, which in Scaloposaurus and Icticephalus connects the lateral and 
posterior pterygoid rami, can be seen. The basisphenoid, however, 
appears to be of the same broad nature as in the Scaloposaurids, and, 
although the specimen is not very well preserved in this region, there 
does appear to be some evidence that the pro-otic is applied to the 
sides of the basisphenoid in a manner very similar to that described 
by Watson in the type of Scaloposaurus. Only part of the epiptery- 
goid is preserved; this is very similar to that of Scylacosaurus and 
Scymnosaurus, 2.e. its dorsal and ventral ends are not expanded, and 
its sides straight; in this character I[ctidosuchoides is thus distinct 
from Trochosaurus and the Whaitsids, and more akin to the generalised 
Pristerognathids. 

In the nature of the dentition, the slight flaring-out of the maxillaries, 
the elongated interpterygoid vacuity, the absence of a postfrontal, the 
slender inferior temporal and postorbital bar, the apparently widened 
basisphenoid, and the large suborbital vacuities bounded by a beam- 
like epipterygoid, Ictidosuchoides approaches the Scaloposaurids. The 
narrow temporal region with its sharp crest, the comparatively long 
and large temporal fossa, the complete postorbital bar, the confine- 
ment of the interparietal to the occipital surface, the large pineal 
foramen, and the apparent absence of a web of bone connecting the 
lateral and posterior pterygoid rami, go to show that Ictidosuchoides 
is a form more akin to the more generalised Pristerognathids, but has 
some parallel characters, which are typical of the Scaloposaurids. 

Chief measurements: 


Length from premaxilla to the basisphenoidal tubera . 140? mm. 
Length from premaxilla to the pineal foramen . 0a 
Length from pineal foramen to edge of occipital plate . 18  ,, 
Length from premaxilla to front of orbit .. 15 ee 


Width across the squamosals «|, 16O2iie 


Morphology of Mammal-like Reptiles of Suborder Therocephalia. 235 


Interorbital width . . ; 24 mm. 
Intertemporal width . ; ; selec lO ee 
Width of the snout . : : : ile wie 
Width across the lateral pterygoid anaes : She Pd Or onl 
Height of the snout . , sa wt hy ag 
Height of the occiput. : : : : wy an28 2 . 
Length of the molar series : ! : P : 2 ies eh 
Diastema ; : : ‘ ie 4 
Length of the three canines. é ‘ 4 18 eee 


Type, B.M.N.H., R5744, Bruintjieshoogte, Somerset Hast, Cape 
Province. 


The family, Lycideopsidae, is instituted for the reception of a single 
crushed skull; this skull cannot be included in any of the existing 
families without affecting the homogeneity of these groups. The 
characters distinguishing this from the other families are—extremely 
long dorso-ventrally depressed skull, with an extremely long molar 
series (10 teeth), consisting of ill-developed teeth, which may be 
absent in old age. 


Lycideops longiceps, Broom. 
(Figs. 15-16.) 

In his original description of 1931, Broom gave little more than the 
dental formula, 1.5, c.1,m.8. In 1932, a slightly fuller description 
and a restoration of the anterior half of the left side was given without 
the constituent elements being indicated. 

Although considerably crushed and sun-cracked, it has been possible 
to clear the external surfaces and most of the palate of a considerable 
amount of matrix. The extensive cracking, however, makes it 
extremely difficult to determine the structure. The dental formula is 
ue 53. 2; mii 10 
mere. m5 
crowded, and in the lower jaw there are no incisors at all; in the upper 
jaw two long and slender canines are present, but it would appear that 
the posterior one is being replaced; in the lower jaw there is only one 
canine, situated very far anteriorly; the upper molars are exceedingly 
small teeth, occupying as much as 50 mm.; they are well separated, 
and appear to be rudimentary; in the lower jaw the molars are also 
very small, with large interspaces; they occupy 26 mm.; there is a 
diastema anterior as well as posterior to the canine. Broom is 


The incisors of the upper jaw are very small and rather 


236 Annals of the South African Museum. 


probably right in thinking that the molars eventually disappear, and 
that Lycideops would in old age be molarless. 

The skull has lost the greater part of its posterior third; on the left 
side, however, the lower jaw is complete and is in articulation with the 
quadrate and quadratojugal; part of the squamosal is also preserved. 

The species is remarkable for its great relative length and con- 
sequent slenderness; the snout, in particular, is very long and slender, 
and was rounded in section; an antorbital depression, if any, must 
have been shallow; the orbit was large and rounded, directed as much 
outwards as upwards; the length of the molar series is very great. As 
far as can be determined, the palate is typically Therocephalian, with 


Ect Peer Poe 


ee 


Fig. 15.—Lycideops longiceps. B.M.N.H., R5695. Lateral view of the 
skull, with the distortion corrected. x 4. 


all the elements relatively lengthened; the transverse pterygoidal bar 
is deep; the suborbital vacuities fairly large; the anterior ramus of 
the pterygoid is very long; the vacuities for the internal nares are 
extremely short, absolutely as well as relatively; they are nearly 
rounded and the anterior part, which received the lower canine, 
situated very far anteriorly. The lower jaw 1s very long and slender, 
and the symphysis is extremely weak; there are no incisors, and the 
splenial, if present, must have been weak, especially at the symphysis. 

This form, with its incipient molar reduction, its depressed form and 
rounded snout, narrow, transverse pterygoidal bar, and the curved 
contour of the mandible, has much in common with the Whaitsids. 
Its great length and the presence of a long molar series, however, 
excludes it from the Whaitsids proper. Lycideops may thus be 
regarded as a form, which in certain respects foreshadows the Whaitsid 


structure. 


Morphology of Mammal-lke Reptiles of Suborder Therocephalia. 237 


Fie. 16.—Lycideops longiceps. B.M.N.H., R5695. 


A. Inner view of the left mandible. x 4. 
B. Outer view of the left mandible. x 4 


Chief measurements : 


Length from premaxilla to front of the orbit . 120 mm. 


Length of the lower jaw : ; 5 ello 
Interorbital width : : : : ; 32a 

Width of the snout. : : : LOW 

Width across the lateral pterygoidal flanges . 58 ,, 
Height of the snout. : : . BO 5, 
Height of the mentum. 4 ; , ; Dias 
Length of the molar series. hee D0: 3 
Diastema . ; ; : 13 or 22 mm. 
Length of the incisor series . 24 mm. 


Type, B.M.N.H., R5695, Thaba ’Nchu, Orange Free State. 


The family, Whaitsidae, is particularly well represented in the 
collection in the British Museum, viz. a good anterior half of the 
type-skull of Moschorhinus kitchingi, a practically complete skull 
of the type of Notosollasia laticeps, the imperfect skull of the type 
of Theriognathus microps, the excellent type-skull, and an additional 
snout, of Whaitsia major, and the imperfect snout, which is the type 
of Tigrisuchus simus. All these forms fall into a homogenous group, 


238 Annals of the South African Museum. 


having all the characters enumerated by Haughton in his original 
description of Whaitsia platyceps. The Whaitsidae may be defined 
as follows: Therocephalians with a broad, depressed, moderately 
short and rounded snout; large temporal openings; crested parietal 
region with a small pineal foramen; molars absent or obsolescent; 
width across the lateral pterygoidal flanges reduced; the anterior 
palatal vacuity greatly shortened, wholly or partly separated into 
an anterior foramen 1ncisivum, and a posterior internal narial opening; 
the anterior part of the palate deeply vaulted with an ingrowing of 
the maxillaries, which is sometimes met by an outwardly directed 
swelling of the prevomers; suborbital vacuity variable, absent, 
small or large; epipterygoid broad, with expanded dorsal and ventral 
ends; postfrontal absent; very strong paroccipital process; weak, 
but long, mandibular symphysis and distinctively curved dentary. 


Moschorhinus kitchingi, Broom. 
(Figs. 17-19.) 

Broom’s original account and figures are good; but, since I have 
developed and etched the skull, some additional features can be 
recorded. Although it is possible that in life six incisors were present, 
this is by no means “quite manifestly”? so; actually, on either side, 
only five are preserved; on both sides there are no functional molars, 
but sockets of two are preserved. The dental formula can thus be 
givenasi.5,c.1,m.2%. Inthe accompanying figures of the dorsal and 
lateral surfaces a number of additional features are shown; it is 
evident that the septomaxilla is a much larger bone than Broom 
thought—its posterior extent, in particular, is very large; the short 
frontals form only a small portion of the supraorbital border; there 
is no postfrontal; the lacrymal carries a distinctive tubercle; there 
is no step in the dentigerous border; the ectopterygoid meets the 
jugal. 

The palate of Moschorhinus is essentially similar to that of the 
other members of the family Whaitsidae, but there are some very 
interesting and illuminating differences in the less basic points of 
structure; the entire palate is relatively wider than in the other 
Whaitsids, and, due to its absolute greater width and less vaulted 
nature, creates an impression of still greater width; this is due to 
the fact that the maxillaries and, to a lesser extent, the palatines do 
not curve inwards as much as they do in the other Whaitsids; in 
Moschorhinus the alveolar edge is fairly straight, whereas in the 
other Whaitsids it sweeps inwards with a medially directed convexity; 


Morphology of Mammal-hke Reptiles of Suborder Therocephalia. 239 


on the inner vertical surface of the palatines there is a sharp oblique 
ridge for the attachment of the soft palate; this is situated half-way 
up, whereas in Notosollasia, Theriognathus, and Whaitsia it lies on 
the ventral edge. 

The prevomers are of a distinctive shape; anteriorly, they are 
greatly expanded, and carry only a very low ventral keel on their 
middle third; a fracture reveals the fact that the premaxillaries 


Jw.— 


Orb, 
Po. 


Fie. 17.—Moschorhinus kitchingi. B.M.N.H., R5698. Dorsal view of the 
: anterior half of the skull. x 4. 


extend on to the dorsal surface of the prevomers; posteriorly, the 
prevomers underlie the pterygoids and palatines and present a shovel- 
shaped outline; they form the greater part of the posterior margin 
of the extremely short internal nares; there is no ventro-lateral 
swelling of the prevomers, which in Notosollasia and Whaitsia 
divides the anterior palatal vacuities in two unequal parts; in 
Moschorhinus these vacuities are short but wide, and slightly con- 
stricted to produce a dumb-bell shape; the whole functioned as 
internal nares, since the lower canine was received in a depression 


240 Annals of the South African Museum. 


Pe, Mx. 


¥, 


OAC ik ee 


£: 


a § <7 
et 


1 


2 
va 
. PAZ 


Fic. 18.—WMoschorhinus kitchingt. B.M.N.H., R5698. Ventral view of the 
anterior half of the skull. x 3. 


PO-b 


Fig. 19.—Moschorhinus kitchingi. B.M.N.H., R5698. Lateral view of the 
skull; posterior half reconstructed. x 4. 


Morphology of Mammal-like Reptiles of Suborder Therocephalia. 241 


in the premaxilla, anterior to the vacuity and the upper canine, 
whereas in Whaztsia the anterior part functioned as a foramen in- 
cisivum, and the posterior as internal nares; the ectopterygoids are 
strongly developed; they form the lateral borders of the large sub- 
orbital vacuities and, posteriorly, send a strong flange to support 
the anterior surface of the transverse pterygoidal bar, but do not 
form the whole corner of this bar as they do in Whattsia, Theriognathus, 
and Notosollasia. The nature of the suborbital vacuities is more 
akin to that of the Scaloposaurids and the less advanced Pristero- 
gnathids, than to that of the other Whaitsids, where they are very 
small or absent; in the Gorgonopsians they are also very small or 
absent. 

The pterygoids are only partly preserved; owing to the size of 
the suborbital vacuities and the posterior extent of the prevomers, 
the pterygoids have, anteriorly, a much smaller palatal surface than 
in the other Whaitsids; as is shown in the figure, a very deep but 
thin median keel is developed on the ventral surface of the anterior 
rami of the pterygoids—a feature not found in the other Whaitsids; 
lateral to this there lies a deep groove, and then there lies the deep, 
thin, postero-medially directed flange of the lateral rami of the ptery- 
goids; there is no interpterygoid vacuity. Unfortunately, nothing is 
preserved of the posterior half of the skull. 

The skull of Moschorhinus kitchingi throws an illuminating light 
on the nature of the Whaitsids. The broad depressed snout, the 
apparently large temporal openings, the reduced width of the lateral 
pterygoidal bar, the absence of the postfrontal, the reduction of the 
anterior palatal vacuities, the slight inward swelling of the maxillaries 
and the palatines, and, finally, the reduction of the molar series, 
clearly stamps it as a Whaitsid. The persistence of two molars, the 
unipartite anterior palatal vacuities, the large suborbital vacuities, 
and the ectopterygoid not forming the corner of the pterygoidal bar, 
show this species to be less advanced than the Whaitsids—Therio- 
gnathus, Notosollasia, and Whaitsia, and probably indicates the 
road traversed by these more advanced forms. In an unnamed 
species of Moschorhinus, described by Broom, there are two per- 
sistent molars in a skull which is mature; and in the inadequately 
known Moschorhinus warreni there are three upper and four lower 
molars. We-thus appear to have a progressive series showing molar 
reduction. 

The relations of Moschorhinus, and thus all the Whaitsids, to the 
other Therocephalians are not very clear; the three specimens of 


242 Annals of the South African Museum. 


Moschorhinus are manifestly considerably removed from the less 

advanced Pristerognathids, and only the discovery of some forms 

less advanced than Moschorhinus will enable one to trace the ancestors 

of the Whaitsids and also of the parallel family of Scaloposaurids. 
Chief measurements : 


Length from premaxilla to pinealforamen . 145 mm. 
Length from premaxilla to front of orbit 93.3 
Interorbital width : : . ; 5G" aes 
Intertemporal width . : : : 1 Onno 
Width of the snout : : Sa ae 
Width across the lateral feeadal Hanon ; oO re 
Height of the snout. 1 DOG 
Length of the molar series . : 4 : ies 
Diastema . : i ; Ae gs 
Length of the incisor series . : A 


Type, B.M.N.H., R5698, Bethesda Road Station, Kea: Province. 


Notosollasia laticeps, Broom. 
(Figs. 20-24.) 

Broom founded this new genus in 1925, and in his description gave 
little more than the dental formula—i. 4,c.1,m.0. In 1932, he gave 
the formula—i. 5, c. 1, m. 0, and stated that a small secondary palate 
is formed by the union of the maxilla and a downward development 
of the prevomers; he could find no suborbital vacuity. 

I have spent a considerable time in removing masses of matrix 
and in etching the skull, which now shows a number of very interesting 
points of structure. 

In the upper jaw there are four well-developed incisors with longi- 
tudinal grooves, but with no serrations on either the anterior or 
posterior edges; these are followed by a very much smaller fifth 
incisor, which appears to be in the process of eruption; between the 
last incisor and the canine there is a space of 4 mm.; the canine is a 
curved tooth of medium size; on the left side there is another canine 
being absorbed, whereas on the right there is a filled-in alveolus 
behind the functional canine; posterior to the canines, there are no 
indications of molars whatsoever. Noftosollasia thus differs in this 
character from Moschorhinus, where two molar alveoli or even three 
teeth persist. 

The palate of Notosollasia is of some interest; anteriorly, the paired 
prevomers meet the palatal processes of the premaxillaries and form 


PO-b. 


-ann 7. 
Snr} 


Fic. 20.—WNotosollasia laticeps. B.M.N.H., R5699. Lateral view of the 
skull. x 3. 


Fic. 21.—Notosollasia laticeps. B.M.N.H., R5699. Ventral view of the 
skulle <a. 


244 Annals of the South African Museum. 


a comparatively narrow internarial bar, which in life supported a 
cartilaginous septum dorsally, and which separates two anterior 


, 
, 


me Bdph. Cu. + On Su, 


Fic. 22.—WNotosollasia laticeps. B.M.N.H., R5699. Occipital view of the 
skull. x 


3 
5° 


fenestrae in the palatal roof, in which the lower canines fit. Posterior 
to the border of these fenestrae the prevomers swell out and become 


Pa. [ Re. 


Fic. 23.—WNotosollasia laticeps. B.M.N.H., R5699. Lateral view of the left 


side of the brain-case, with the occipital plate seen in section. x 2. 


thickened ventrally as well as laterally, so that they meet the dorso- 
medially bulging maxillaries; the ventral surface of the prevomers 
still, however, occupies a plane much dorsal to the alveolar maxillary 
border. Continuing in posterior direction the prevomers again 
become narrow, and lateral to them lie the two internal nares; the 


Morphology of Mammal-like Reptiles of Suborder Therocephalia. 245 


prevomers curve round the posterior border of the internal nares 
and then continue in posterior direction to meet the palatines and 
the pterygoids, as shown in the figure. The posterior two-thirds of 
the prevomers carry a high keel on the palatal surface, and on their 
dorsal surface are supported by a girder formed by the anterior 
pterygoidal rami (this is clearly shown in a cross-section). The 
maxillaries curve downwards and inwards to a great extent, and, 
in addition, are flanked by the palatines, so that the prevomers form 
the roof of a very narrow, grooved naso-phalangeal passage, which 
lies at a much higher level than the surface of the antero-lateral 
portions of the palatines. The relations of the pterygoids, palatines, 
and ectopterygoids are shown in the figure and are basically as in 
the less advanced Pristerognathids. The small size of the rounded 
suborbital vacuities is a feature which appears typical of the Whaitsids 
(with the exception of Moschorhinus), and is in marked contrast to 
the other Therocephalian families, where it is always large. There 
are no teeth on the palatines and pterygoids, which is in accordance 
with the general lack of tooth-development in the Whaitsids. Pos- 
teriorly the pterygoid sends out a process, which eventually abuts 
on the quadrate, and, in the median line, two high flanges form a 
strong keel, which separate to clasp the anterior prolongation of the 
basisphenoid,. which apparently represents a parasphenoid fused 
with the basisphenoid proper. 

The brain-case and basicranial region have been adequately freed 
of matrix. The brain-cavity is relatively long, narrow, and low. 

The basioccipital forms a strong condyle, and sends two strong 
ventrally directed processes to support the basisphenoidal tubera; 
these processes form the posterior border of the foramen ovale; below 
the level of the foramen magnum there is thus a considerable extent 
of basioccipital. 

The basisphenoid is not very strongly developed; its tubera are 
unthickened, and they do not diverge much; dorsally, it meets the 
pro-otic and forms the posterior border of the lateral opening into 
the pituitary fossa; unfortunately, the relations of the anterior 
prolongation of the basisphenoid cannot be determined; on the lateral 
surface of the vertical sheet there are two horizontally placed basi- 
pterygoid processes clasped by the pterygoid. 

The pro-otic is fairly well developed; antero-dorsally of the fenestra 
ovalis there lies a small foramen for the seventh nerve; from here 
a ridge runs anteriorly to the anterior pro-otic process; above this 
lies the large perforation for the fifth nerve; dorsal to this lies the 


246 Annals of the South African Museum. 


posterior pro-otic process, which forms the border of the large venous 
foramen; anteriorly, the pro-otic is flanked by the broad epipterygoid, 
and, posteriorly, is applied to the paroccipital and supraoccipital. 
Dorsal to the pro-otic the side-wall of the posterior part of the brain- 
case is formed by flanges from the interparietal and parietal. 


Fic. 24.—Notosollasia laticeps. B.M.N.H., R5699. 


A. Outer view of the mandible. x 
B. Inner view of the mandible. x 


The epipterygoid is a low, broad bone with greatly expanded ends; 
it is firmly attached to the quadrate ramus of the pterygoid ventrally, 
and, dorsally, to the parietal; it encloses a large cavum epiptericum. 

The greater part of the occiput is preserved; it is low and broad; 
the post-temporal fenestrae are large and situated low down (contrast 
Trochosaurus); the foramen magnum is small, and attention must 


be drawn to the great depth of bone ventral to it; the paroccipital 


bar is strong, and lateral to it the squamosal forms a deep auditory 


Morphology of Mammal-like Reptiles of Suborder Therocephalia. 247 


groove; the exoccipital takes no part in the formation of the condyle, 
but forms a strong tubercle overhanging the large foramen jugulare: 
the quadrate and quadratojugal are intimately connected, and the 
latter is exposed on the lateral surface; the stapes is preserved on the 
right side in its natural relations; it is a strong bone with expanded 
ends and constricted shaft; no foramen can be seen. 

The lower jaw is of a distinctive shape when viewed from below; 
at its articulatory end the ramus is very wide; it then sweeps forward 
in a medially directed curve and, at the level of the canines, it 
straightens out and forms a long and relatively narrow symphysis. 
It has not been possible to expose the extreme tip of the ramus; 
a fracture near the end, however, reveals a large canine, oval in cross- 
section, and another in the process of eruption; no roots of incisors 
are visible at this plane, and it does not appear probable that any 
could be present, as the canines do not leave much room anteriorly 
for the development of additional teeth; there are no molars. The 
general arrangement of the constituent bones are typically Thero- 
cephalian; their relations are best understood by referring to the 
accompanying figure (fig. 24). 

Chief measurements : 


Length from premaxilla to basioccipital condyle 210 mm. 


Length from premaxilla to front of orbit eh LOOM! 
Length of the lower jaw : : oo) Sie 
Width across the squamosals ; ~. - oS, 
Width of the snout. : : OOme. 
Width across the quadrates . ; eee OES 
Width across the lateral pterygoid fampes GOs, 
Height of the snout. ; : : : A 
Height of mentum of lower jaw . tae, 
Length of the incisor series . : : 7s a 


Type, B.M.N.H., R5699, Bethesda Road Station, Cape Province. 


Theriognathus microps, Broom. 
(Figs. 25-27.) 

Owen’s figures give a good idea of the skull as it was preserved. 
This author considered the specimen to represent an Anomodont, 
and Lydekker, subsequently, placed it in the genus Hndothiodon. 
Broom, in 1910, gave a good summary of Owen’s and Lydekker’s 
accounts and, on a superficial study of the lower jaw, rightly came 


248 Annals of the South African Museum. 


to the conclusion that it was a Therocephalian. In 1932, Broom 
thought it best to regard it as a member of the Whaitsidae. 

Mr. Parsons has attempted to expose the palate, but found that 
for the greater part he could not remove the matrix without damaging 


Fic. 25.—Theriognathus microps. B.M.N.H., 47065. Ventral view of the 
skull. x 4. 


the bone. Subsequently, with a hammer and a small chisel, I 
succeeded in exposing the whole palate and also parts of the occiput. 
A fortunate fracture exposed the outer surface of the left epipterygoid 
and proved that the matrix would split very easily. I thus deliber- 
ately induced additional fractures, with the result that the entire 
brain-case can now be studied from a variety of sections—frontal, 


Morphology of Mammal-like Reptiles of Suborder Therocephalia. 249 


sagittal, parasagittal, and cross-sections—and also from some parts 


where it has been possible to expose the outer surface. 


, 


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B.M.N.H., 47065. 


Fie. 26.—Theriognathus microps. 
A. Sagittal section, reconstructed from a number of partial frontal, parasagittal, 
sagittal, and cross-sections. x 4. 
B. Cross-section through the brain-case, just anterior to the pituitary fossa. x 1. 


22 


It is now clear that, basing his conclusions mainly on the molarless 
nature of the jaws and the structure of the mandible, Broom was 


VOL. XXXI, PART 2. 


250 Annals of the South African Museum. 


right in referring this form to the family Whaztsidae. The general 
appearance and arrangement of the various palatal and occipital 
bones and the structure of the brain-case bear a very close 
resemblance to that of Whaitsia, Moschorhinus, and Notosollasia. 

The anterior end of the snout is missing and nothing can be ascer- 
tained as to the nature of the canines and incisors; as both alveolar 
edges have been perfectly exposed it is possible to state definitely 
that no molars were present; there are not even indications of alveoli. 
The dorsal and lateral surfaces have for their greater part been lost, 
and the occipital surface has also been much damaged. 

The palatal surface, although somewhat sun-cracked, is preserved 
in a sufficiently complete state for its whole structure to be deter- 
mined. The relations of the constituent elements are shown in the 
accompanying figure. Although only the posterior borders of the 
anterior palatal vacuities are preserved, it would appear that they 
were of the type of those in Moschorhinus rather than that of Notosol- 
lasia and Whaitsia, 2.e. the whole functioned as internal nares, the 
canines of the lower jaw being received in hollows antero-laterally 
to the nares. The prevomers are relatively long and form the 
median part of the roof of the deeply vaulted anterior half of the 
palate. On both the dorsal and ventral surfaces the prevomers 
carry high median keels. Lateral to the prevomers he the palatines, 
which form the deep lateral walls to the highly vaulted median part 
of the palate. The palatines extend laterally to meet the maxillaries 
at the edentulous alveolar edge; on their medial surface the palatines 
carry a well-defined ridge, to which in life the soft palate must have 
been attached; between the prevomers and the soft palate a spacious 
air-passage was thus developed. The ectopterygoids are remarkable 
in that they form the strong lateral corner of the transverse pterygoidal 
bar, whereas, in the normal Therocephalian condition, and also in Mos- 
chorhinus, they are only supporting flanges to the pterygoids, which 
themselves form the corners. A small suborbital vacuity is present; 
this is very similar to that of Notosollasia, whereas it is large in 
Moschorhinus and, by both Broom and Haughton, maintained to be 
absent in Whaitsia platyceps, as it isin Whaitsia major. The pterygoid 
is very well developed; it forms a large part of the middle portion 
of the palate; the lateral flanges are not very strong since the corners 
of the pterygoidal bar are formed by the ectopterygoids; the posterior 
rami form, in their anterior part, two diverging flanges, in between 
which lies a deep groove; continuing posteriorly, these flanges form 
a very deep keel and clasp between them a similar high unpaired keel 


Morphology of Mammal-like Reptiles of Suborder Therocephalia. 251 


sent forwards by the basisphenoid (basisphenoidal rostrum); the 
quadrate rami stretch postero-laterally with a straight outer edge 
(as is typical for the Therocephalia in contradistinction to the 
Gorgonopsians, where it is curved) to meet the quadrate. 

The basioccipital (figs. 25-27) forms the well-developed condyle, 
and, in a fracture, the processes supporting the basisphenoidal 
tubera are very well shown; dorso-laterally the exoccipitals support 
the condyle; the medulla rested on its rather irregular grooved 
dorsal surface. 

The basisphenoid forms two widely separated tubera and, 
anteriorly, forms the deep vertical keel flanked by the pterygoids; 
the basipterygoid processes 
are horizontal, flat, and 
immovable; in a sagittal 
fracture no trace ofa median 
parasphenoidal process is 
seen extending dorsally, 
although both the anterior 
and posterior borders of the 
pituitary fossa can be seen; 
it is, however, possible that ais 
the anterior partofthe bone Fie. 27.—Theriognathus microps. B.M.N.H., 
labelled “ basisphenoid” re- eee Lateral nay of the left side of the 

; rain-case. x #4. 
presents a parasphenoid 
indistinguishably fused with it; in a cross-section it is seen that the 
basisphenoid forms a broad floor for the hind-brain. 

The pro-otic is only partly exposed and, as far as can be ascertained, 
agrees with the condition in Notosollasia; it lies against the par- 
occipital; antero-dorsally, it forms the ventral border of the venous 
foramen; the foramen for the seventh nerve apparently lies in the 
usual position; that for the fifth is obscured by the epipterygoid. 

The epipterygoid, as in all the Whaitsids, is a large flat element 
with expanded dorsal and ventral ends; its base on the quadrate 
ramus of the pterygoid is very long and, as it lies lateral to the cavum 
eprptericum, obscures the foramen for the fifth nerve, whose nature 
has thus not been determined. 

The occiput is depressed; the post-temporal fenestrae are of medium 
size; the paroccipitals are very massive, with greatly expanded distal 
ends, which support the auditory ridge formed by the squamosals; 
they are situated in a plane considerably in advance of the par- 
occipital processes. 


252 Annals of the South African Museum. 


The lower jaw is of the usual Therocephalian structure, but in 
shape is curved in the distinctive way seen more highly developed 
in Notosollasia and Whaitsia. The angular is a strong element with 
a large exposure externally; the surangular forms the usual curved 
girder intercalated between the dentary and the articular; the 
dentary bears no molars; the articular articulates with both quadrate 
and quadratojugal. 

A long slender bone, which is probably a ceratohyal, is imbedded 
in the matrix lying on the right side of the basisphenoidal keel. 

Chief measurements : 


Length from premaxilla to the basioccipital 

condyle . ‘ : . 205? mm. 
Length from eens to fhe Biviea foramen 150? ,, 
Length from pineal foramen to the occipital 


plate : ‘ 30: 
Length from premaxila to “the font a he 

orbit : : : : ; . . 90%ee 
Length of the lower jaw ; . 200 Far 
Width across the squamosals ‘ : Rc ® 
Interorbital width t : ; 5 , aoe 
Intertemporal width . . » AO sae 
Width across the lateral nicreeidal fanees 80) 25; 
Width of the snout. : ; . [igs 
Height of the occiput . 2 : .. 45-7 


Type, B.M.N.H., 47065, Stylkrantz, Sneeuwberg, Cape Province. 


Whartsia major, Broom. 
(Figs. 28-34.) 

Five good skulls of this species are known. The type in the British 
Museum consists of a very good skull, which is, however, slightly 
distorted; associated are a right radius, ulna, and manus; the latter 
was described by Broom as a pes; a humerus is catalogued (R5755) 
as that of Whaitsia, and the nature of the matrix and of the fossilisa- 
tion is as in the skull; the femur described by Broom is not in the 
British Museum collection; a snout labelled as Anteosaurus minor 
is obviously that of another specimen of this species: its number is 
R5748. 

Through the courtesy of Dr. W. E. Swinton, Mr. Parsons was asked 
to do the preliminary clearing of the type skull; I have carried the 


— EEE = 


Morphology of Mammal-like Reptiles of Suborder Therocephalia. 253 


Fic. 28.—Whaitsia major. B.M.N.H., R5694. Dorsal view of the skull. x 33. 


depp Dent. 


Fic. 29.—Whaitsia major. B.M.N.H., R5694. Lateral view of the skull. x j- 


254 Annals of the South African Museum. 


development further so that it now shows the greater part of the 
detailed structure. 

The skull is of a large Therocephalian; it is broad and depressed; 
the intersquamosal width is great and the temporal fossae are, as in 
all the Whaitsids, thus very wide, but short; the posterior part of 
the mandibular ramus 1s very wide, but in anterior direction it curves 
inwards and forms a narrow, though long, symphysis. 

The dental formula is 1. 5, ce. 1, m.0; asin Notosollasia the incisors 
carry longitudinal grooves. 

The structure of the outer surface is best understood by referring 
to the figures of the dorsal and lateral surfaces; the parietal crest 
is thin and fairly high; the pineal foramen is a narrow slit; the 
frontal is broad, but short, with only a small entry into the supra- 
orbital border; there is no postfrontal; the dorsal and anterior 
borders of the orbit are raised into a sharp ridge, which is interrupted 
above the lacrimo-prefrontal suture; the lacrimal and prefrontal 
form a deep anterior face to the orbit; there is a small lacrimal 
foramen; the facial exposure of the septomaxilla is fairly large, and 
it has two distinctive spurs projecting into the nostril; the two 
depressions along the maxillo-septomaxillary suture, usually found 
in Therocephalians, are very well shown. 

The occiput is exposed exceedingly well; the basioccipital condyle 
is strong and kidney-shaped; it sends down two strong processes to 
support the basisphenoidal tubera; the exoccipitals support its 
corners without actually coming on to the articulatory surface, and 
form quite strong processes overhanging the large jugular foramina; 
the paroccipital processes are strong; the post-temporal fenestrae 
are of medium size, and are situated low down; the foramen magnum 
is small; the tabular is a large bone forming a considerable part of 
the post-temporal edge; the quadrate and quadratojugal le well 
below the level of the condyle. 

The palate has its posterior half well exposed in the type, where 
it is, however, distorted; the anterior half is beautifully shown in 
the duplicate specimen—R5748, and the accompanying figure is a 
composite one based on both specimens. In the anterior half the 
important point to notice is the bipartite anterior fenestra; the 
anterior part received the lower canine, and the posterior functioned 
as internal nares, posterior to which the palatines form a deep naso- 
phalangeal passage; the partition is brought about by a process of 
the maxilla growing in dorso-median direction and then overlying 
a ventro-laterally directed process sent out by the prevomer. In 


Morphology of Mammal-like Reptiles of Suborder Therocephalia. 255 


Fig. 30.—Whaitsia major. B.M.N.H., R5694. Ventral view of the skull. x 45. 


} eres | iH 
an ) 


Pe X BOc. BNph. Exe St 


Fic. 31.—Whaitsia major. B.M.N.H., R5694. Occipital view of the 
glaullls 8 Bs 


7 


256 Annals of the South African Museum. 


Moschorhinus the swelling of the maxilla inwards has been mentioned, 
but here there is no outgrowth from the prevomer; in Notosollasia 
the maxilla grows inwards and the prevomer has a lateral process, 
which meets it so that the vacuity is bipartite, but here the process 
of the prevomer is in a plane considerably dorsal to the maxillary 
swelling; in Whzitsia the maxillary and prevomerine processes lie 
in the same plane, which les at an angle of about 45° to the horizontal. 
In this character these three forms form a morphological series, which 
is, however, certainly not a direct ancestral one. Posterior to the 


. ie 
<7 


aA\e SE Per — PSph. 


Fit. 


ies 
Fie. 32.—Whaitsia major. B.M.N.H., R5694. Lateral view of the right 
side of the brain-case. x 3. 


internal nares, the prevomers expand in shovel-shaped fashion and 
underlie the palatines and pterygoids. On the anterior ramus of 
the pterygoid there are two large and deep depressions; I can offer 
no suggestion as to their significance. There is no suborbital vacuity. 
The ectopterygoids, as in Notosollasia and Theriognathus, form the 
corners of the transverse pterygoidal bar, whereas in Moschorhinus 
it is still formed by the pterygoid itself. Posteriorly, the pterygoids 
form a deep keel in the median line, and here clasp the anterior pro- 
longation of the basisphenoid, which here may have a parasphenoid 
fused on to its ventral surface. The quadrate ramus has a straight 
outer edge and is applied to the anterior face of the quadrate. The 


Morphology of Mammal-lke Reptiles of Suborder Therocephalia. 257 


basisphenoid is not a very strong bone and its tubera are, for their 
greater part, formed by the overlying basioccipital processes; it is, 
however, possible that the bone forming the ventral surface of the 
tubera is really a parasphenoid, and that the processes supporting 
it are really basisphenoidal, and not basioccipital; the sagittal fracture 
in Theriognathus, however, shows a good suture between the basi- 
occipital and basisphenoid, with nothing to indicate that the ventral 
part of the basisphenoid is really an underlying parasphenoid very 
closely applied to the basisphenoid proper. In ventral view it is 
seen that the basisphenoid swells out and meets the pro-otic, which 
then continues dorsally. The quadrate forms the main articulatory 


Spl cart PB Are. Ang 


Fic. 33.—Whaittsia major. B.M.N.H., R5694. Inner view of the right 
mandible. x 4. 


surface, and the quadratojugal only the lateral corner, which, in 
lateral view, projects below the level of the squamosals. 

The brain-case is beautifully shown in lateral view on the right 
side; the epipterygoid is a large bone with greatly expanded ends 
and a constricted waist; it lies in a plane lateral to the pro-otic, and 
encloses a large cavum epiptericum; the pro-otic is well developed; 
ventrally, it meets the basisphenoid in a suture, which runs from the 
border of the fenestra ovalis to the lateral border of the pituitary 
fossa; dorsally, it meets a flange of the interparietal and, posteriorly, 
is applied to the paroccipital; it has a strong anterior process, which 
separates the lateral opening into the pituitary fossa from the opening 
for the fifth nerve; it forms part of the anterior border of the fenestra 
ovalis and, antero-dorsally to this, is pierced by a foramen for the 
seventh nerve; a posterior pro-otic process separates the foramen 
for the fifth nerve from the large venousforamen. From this structure 


258 Annals of the South African Museum. 


of the lateral wall it is clear that the posterior part of the brain-case 
is fairly wide and not very high, with a mass of bone below its ventral 
A floor; its anterior extent is 

not great, and it housed 

only the hind-brain; the 
mid-brain and thalamen- 
cephalon lay median to the 
epipterygoid, and the whole 
cerebrum must have been 
housed by the ethmoidal 
part, which, in this speci- 
men, has not been exposed. 
The lower jaw is char- 
acterised by the great width 
across the condyles and 
then, anteriorly, by the in- 
wardly curved sweep to the 


long and narrow symphysis; 
there do not appear to be 
any incisors, and there are 
no indications of molars 
whatsoever; the splenial is 
very intimately fused to the 
dentary; the coronoid is 
small, and the surangular 
strong; the angular is large, 
but extremely thin. 

ee The fore-limb is repre- 
“sented by a good right 
humerus, which may be of 
the same animal as the 
right ulna, radius, and 
manus. The humerus is of 


Fic. 34.—Whaitsia major. a type distinct from that 
A. Dorsal view of the right fore-foot. known in any other Therap- 
B.M.N.H., R5694. x 3. sid; there is a very large 
cl—c3 =centrals. Ua tilnar : 
is =intermedium. ws =ulnare. Eons icondy lar, but no 
Ro) radius: 1-5. =distalia. ectepicondylar foramen; 
r. =radiale. I-V. =digits. 


the delto-pectoral crest is 
very long; distally, it merges into the shaft; it curves inwards and 
does not lie along the anterior border as in the Gorgonopsians, e.g. 


Morphology of Mammal-like Reptiles of Suborder Therocephalia. 259 


Aelurognathus, Arctognathoides, etc.; the surface for the insertion of 
the deltoid muscle lies less on the dorsal surface than it does in 
the Gorgonopsians and Anomodonts; the distal end is greatly ex- 
panded, the ectepicondyle being particularly strong; the scar of the 
scapulo-humeralis posterior and subcoraco-scapularis is very strong, 
and so is the supinator and extensor crest; the large pit for the 


Fic. 34.—Whaittsia major. 


B. Ventral view of the right humerus. B.M.N.H., R5755. x 
C. Dorsal view of the right humerus. B.M.N.H., R5755. x 


ent. =entepicondyle. 


to be 
Ser 


brachialis indicates a strong muscle of that name; the intertrochanteric 
fossa 1s deep, and the coraco-brachialis must have been strong. 

The ulna is a very peculiar bone; there is no olecranon process, 
but the proximal end is greatly expanded preaxially in a manner 
unknown in any other Therapsid; the bone is somewhat flattened, 
and this expansion may represent a sigmoid process with a sigmoid 
notch into which the radius fitted. 

The radius has no distinctive characters; it is slightly shorter than 


260 Annals of the South African Museum. 


the ulna; as the humerus is also short, the whole fore-limb was short, 
and the animal could thus not have been very agile. 

The carpus is not fully preserved, the intermedium and the greater 
part of the ulnare being lost; the radiale is a strong, oval-shaped bone, 
to which, distally, two bones are articulated; the preaxial one is 
undoubtedly a composite bone consisting of the fused first and second 
centralia; the central one is the third central; there are four distals; 
the postaxial one, which articulates with the fourth and fifth meta- 
carpals, represents the fused fourth and fifth distals; the meta- 
carpals have peculiar mammal-like epiphysial ends, also encountered 
in the Gorgonopsian, Aelurognathus; the phalanges have very well- 
modelled articulatory surfaces with a much better finish than those 
of Aelurognathus. The carpal formula is thus 3, 3, 4, and the phal- 
angeal 2, 3, 3, 3, 3, as in Cynodonts and primitive Mammals, and not 
2, 3, 4, 4, 3, as in Aelurognathus. 

Chief skull measurements : 


Length from premaxilla to the basioccipital condyle . 305 mm. 
Length from premaxilla to the pineal foramen A .. BOOT 
Length from premaxilla to the occipital plate. : Ue 
Length from premaxilla to the front of the orbit . -: PROT 
Length of the lower jaw : : ~ Zoom 
Width across the squamosals : ; > 4 2p0' a 
Interorbital width : : : . : . : 65-5 
Intertemporal width . : : : ; : ; 1: = 
Width of the snout. : : , SD os 
Width across the lateral Meee oid dunes : 507 
Height of the snout . : : : ‘ F ; ro 6 hae 
Height of the occiput . : : : Ue 
Length of the incisor series . : Say 


Type, B.M.N.H., R5694, Thaba ’Nehu, One cene Free State. 


Tigrisuchus siumus, Owen. 


This very unsatisfactory type consists of a weathered snout, which 
shows the roots of three large incisors and a large canine on both sides 
of the upperjaw. This snout has always been considered to represent 
a species of Gorgonopsian, but recently Broom has recognised it to 
be a Therocephalian. In comparing it to the Whaitsid skulls in the 
collection, I was impressed by the similarity in the broad depressed 
snout, the general shape of the nostril and the septomaxillary, and 
the broad expanded anterior end of the prevomers. As there appears 


Morphology of Mammal-like Reptiles of Suborder Therocephalia. 261 


to be no character which would exclude it from the Whaitsid family, 
this fragment had better be included in this group. The specimen 
is of some morphological interest in that a fracture shows the detailed 
relations of the premaxilla, septomaxilla, and the prevomers. 


Type, B.M.N.H., R1721, Stylkrantz, Graaff-Reinet 2, Cape Province. 


The family Huchambersidae is instituted for the remarkable skull 
described by Broom as Euchambersia mirabilis. The possession of 
the following characters clearly distinguish this new family from the 
other Therocephalian families, viz. jugal bar absent; incomplete 
postorbital bar; the presence of a very remarkable preorbital de- 
pression, which leads by a notch into the buccal cavity; the canine 
has a sharp ridge on its labial surface; there is no maxillary border 
posterior on the canine, and thus there are no molars. 


Euchambersia mirabilis, Broom. 
(Fig. 35.) 


This remarkable skull was first described by Broom in 1931, and 
figured in 1932. The skull, though crushed in an oblique direction, 
is fairly completely preserved; the squamosals, quadratojugals and 
quadrates have been lost, and the occiput cannot be satisfactorily 
cleared. 

The skull is unlike that of any other Therapsid, but the essential 
points of structure in the palate, dorsal and lateral surfaces con- 
clusively show that the animal is an aberrant Therocephalian. There 
are a number of remarkable specialisations, viz. the postorbital and 
infraorbital bars are incomplete; the infratemporal bar is absent, 
though it is likely that the squamosal formed an incomplete bar; 
there is a very large and deep preorbital depression lying in the 
external surface of the maxilla and the lacrymal; a notch leads from 
this cavity to the buccal cavity; there are no molars; the canine 
is of medium size, and has a unique ridge on its labial surface; no 
incisors are preserved, but, according to Broom, five were present. 
Broom has suggested that the preorbital depression housed a huge 
par-otic gland, and Nopcsa suggested that the gland was poisonous, 
and was connected with the ridge on the canine. Broom’s suggestion 
is untenable, as the depression is preorbital, and the par-otic gland is 
generally postorbitally situated. It seems more likely that this 
depression housed an enlarged labial or lacrymal gland. At this 


fen. Ov 


Fie. 35.—Euchambersia mirabilis. B.M.N.H., R5696. x 2. 


A. Dorsal view of the skull. 
B. Ventral e 
C. Lateral a 


29 


29 


Morphology of Mammal-like Reptiles of Suborder Therocephalia. 263 


point it is of interest to recall the fact that in the Gorgonopsians 
and Therocephalians there is generally a shallow preorbital depres- 
sion, which probably housed a gland of some kind; the gland in 
Euchambersia may be the result of the hypertrophy of such a normal 
supralabial or lacrymal gland. 

On the outer surface the important points are: the very large 
prefrontal, which forms the whole supraorbital border; the small 
frontal with no entry into the orbital border; the absence of a post- 
frontal; a minute pineal foramen; the rudimentary jugal forming 
a little of the preorbital border, but not extending backwards to 
meet the squamosal or postorbital; the short maxillary border, 
which does not meet the jugal; the large septomaxilla. 

The palate is essentially Therocephalian; the anterior palatal 
vacuities are unipartite and relatively long; the prevomers are, 
anteriorly, greatly expanded and, posteriorly, are shovel-shaped, 
and underlie the pterygoids and palatines; the ectopterygoid does 
not reach the corner of the lateral pterygoidal bar; the median 
vertical flanges of the posterior pterygoid ramus do not meet along 
the median line, and laterally flank the basisphenoidal rostrum; if 
my interpretation, that the basioccipital sends two strong processes 
ventrally, is correct, then the basisphenoid is a thin bone underlying 
these processes; on the other hand, if these processes are actually 
the basisphenoidal tubera, then the thin bone must be a parasphenoid ; 
I find no evidence for the latter view; the paroccipital bar is strong, 
and is distally grooved to receive the quadrate; dorsal to this the 
supraoccipital sends a thickened process laterally, and to this a 
tabular must have been attached; actually, no tabular is, however, 
preserved; the basioccipital condyle is fairly strong. 

The epipterygoid is a moderately slender element, as in the 
Pristerognathids, Scylacosaurus, and Scymnosaurus; the pro-otic 
is not sufficiently exposed to merit a description; the fenestra ovalis 
is large; there is a great depth of bone ventral to the foramen 
magnum. 

Euchambersia is thus a Therocephalian, which can be derived from 
an unspecialised Pristerognathid; its evolutionary direction has been 
along a line quite distinct from that of the Lycideopsids, Whaitsids, 
and Scaloposaurids; in the main its palate is quite primitive, and so 
is the epipterygoid; but the extreme specialisation in the occiput, 
the loss of the postorbital and jugal bar, the development of the 
peculiar preorbital depression, and the apparently correlated notch 
leading on to the ridged canine, are all characters with no parallel 


264 Annals of the South African Museum. 


in any of the other Therocephalian families. It is of interest to 
recall that an analogous loss of postorbital and infratemporal bar 
connected with the development of a poison fang occurs in the 
Ophidia. 


Chief measurements : 


Length from the premaxilla to the basioccipital condyle . 120 mm. 


Length from the premaxilla to the front of the orbit . | Sao 
Interorbital width : : : : ; . 28a 
Intertemporal width . : : : : . ieee 
Width of the snout : ; : i ae 
Width across the lateral sae antes . ee 
Height of the snout. : ee 
Height of the occiput . : : :  oaees 
Depth of bone below the foramen magnum . : Zee 


Type, B.M.N.H., R5696, Norval’s Pont, Cape Province. 


In the family Scaloposauridae seven genera have been included, 
viz. Akidnognathus, Choerosaurus, Icticephalus, Ictidognathus, Ictido- 
stoma, Scaloposaurus, and Simorhinella. Three of these are repre- 
sented in the collection of the British Museum, and they have recently 
formed the basis of an excellent description by Watson; with the 
exception of a few supplementary remarks on Icticephalus, it is thus 
unnecessary to go into any details. 

The family is of considerable interest in that it ranges throughout 
Lower Beaufort times without showing any very marked evolutionary 
changes; Icticephalus of the Tapinocephalus zone may even be 
cogeneric with Scaloposaurus of the Custecephalus zone. Although 
occurring as early as the Tapinocephalus zone, the Scaloposaurids 
are, in a number of points of structure, definitely distinct from any 
of the other Therocephalian families. The Whaitsids, Lycideopsids, 
and Euchambersids represent three lines of development which are 
completely divergent from that of the Scaloposaurids. The Pris- 
terognathids are apparently a retarded stock in which a number of 
primitive features are retained; in one form, Scylacosaurus, the 
nature of the basisphenoid suggests a development parallel to that 
of the Scaloposaurids. In the Ictidosuchids, Ictidosuchoides has 
been shown to possess some characters also met with in the Scalopo- 
saurids. It would thus appear that the Therocephalians of the Lower 
Beaufort represent the end branches of a phylogenetic tree, whose 
main stem must be sought for in pre-Beaufort rocks. 


Morphology of Mammal-lhke Reptiles of Suborder Therocephalia. 265 


Icticephalus polycynodon, Broom. 


In his original description Broom made the following observations : 
dental formula, 1.6, c.3,m.11; all the teeth are unserrated; this new 
type differs from Scaloposaurus in having eleven, and not nine, molars, 
and in that the postorbital arch meets the jugal. 

Watson, in 1931, gave a much fuller account, and with this I am 
in agreement. I have, however, found that something more can 
be determined of the dentition. Although very badly preserved, 
Broom is probably right in assuming the incisors to be six in number; 
there is a large canine, with one tooth anterior and another posterior 
to it—all on the maxilla; then there is a diastema of 2 mm.; this is 
followed by nine molars, although between the fifth and sixth there 
is an interspace, which may have housed another tooth; the molars 
are small and progressively decrease in size in posterior direction. 
In the British Museum specimen the dental formula would thus be, 
i. 6, c. 3, m. 9-10, which is thus not very different from that of Scalopo- 
saurus. If Broom’s supposed distinguishing character in the dentition 
is thus invalid, the only other distinguishing character is the nature 
of the postorbital bar, and in neither of these two specimens is the 
nature of the postorbital bar known with absolute certainty. Watson 
has at great length drawn attention to the great similarity of these 
two skulls in nearly all, even the most detailed, points of structure. 
So that one can really only maintain that the type Icticephalus of 
the Tapinocephalus zone is represented in the Cistecephalus zone by 
a practically identical animal, Scaloposaurus. The older Icticephalus 
has a slightly larger skull, with its snout higher and thus relatively 
less broad; the orbits look as much outwards as upwards; the 
mentum of the lower jaw is fairly deep. In the younger Scalopo- 
saurus the skull is somewhat smaller; the whole skull, including 
the snout, is somewhat flattened, so that the snout is relatively 
broader than high; the orbits look more upwards than outwards; 
the mentum of the lower jaw is less deep and more sloping. Viewed 
from in front, Scaloposaurus has a more pointed snout and appears 
rounded in section, whereas in Icticephalus it is squarish. 


Type, B.M.N.H., R4096, Weltevreden, Prince Albert, Cape Province. 


Chief measurements: 


Scaloposaurus. Icticephalus. 
Premaxilla to basisphenoid . 50% mm. 65 mm. 
Premaxilla to pineal foramen . a D9 +55 


VO “xX. PART 2. 23 


266 _ Annals of the South African Museum. 


Scaloposaurus. Icticephalus. 
Premaxilla to front of orbit . 282% mm. 34 mm. 
Interorbital width . Eo OES ae IZ er 
Intertemporal width oT cee ee 
Width of the snout . isis 16.49 
Width across lateral ieereoid 
flanges. : re OTs oe 2 
Height of the Laois Kees 83 Lo 
Height of mentum of lowerjaw. 7? ,, Ow 
Length of the molar series : ee Lane 
SUMMARY. 


1. The palate is figured and described in three Pristerognathids, 
one Ictidosuchid, four Whaitsids, and one Euchambersid; it can thus 
be claimed that the main morphological features of this region in 
the Therocephalians have been established. In the more generalised 
Pristerognathids, in the slightly more advanced Ictidosuchids, and 
in the specialised Euchambersids and Scaloposaurids, the palate is 
comparable in general plan to that of the less advanced Deino- 
cephalians and Pelycosaurs; even in the very specialised Whaitsids, 
the underlying ground plan is still the same. The Therocephalian 
palate differs from that of the Gorgonopsians, which occupy the 
same developmental niveau, but on a parallel plane, in the number 
of points mentioned; the chief being the nature of the prevomers. 

2. The occiput is described and figured in one Pristerognathid and 
in two Whaitsids; it differs mainly from that of the Gorgonopsians 
in the very massive paroccipital bar and the large post-temporal 
fenestra (except in Scymnosaurus); in the Scaloposaurids a special 
process is developed on the paroccipital. 

3. Parts of the brain-case are figured and described in two Pristero- 
gnathids, in one Euchambersid, and in three Whaitsids; it has 
become evident that in the Therocephalians there has, in respect of 
the epipterygoid, occurred a development, which has in some respects 
been paralleled by the Cynodonts. 

4. The description of a complete fore-limb of Whaitsia adds to our 
knowledge of the postcranial skeleton of the Therocephalians, about 
which lamentably little is as yet known. 

5. In the course of this paper it has been shown that the seven 
families, into which the suborder has been divided, represent distinct 
evolutionary trends, and that certain similarities in structure do not 
necessarily imply any close direct relationship, but may simply have 


Morphology of Mammal-like Reptiles of Suborder Therocephalia. 267 


resulted as parallel developments from an original stock possess- 
ing common characters and_ potentialities. The Whaitsids, 
Kuchambersids, and Lycideopsids are families possessing some very 
-definite characters, separating them from each other and from the 
Scaloposaurids, Alopecopsids, Ictidosuchids, and Pristerognathids; 
the Scaloposaurids are also quite distinct, but may be a composite 
family, as there is a discrepancy in one character, viz. in some the 
postfrontal is present, whereas in others it is absent; finally, the 
Pristerognathids, although the most generalised group, does contain 
some forms which possess quite advanced characters, e.g. the 
epipterygoid of Trochosaurus. Some subdivision, as, for example, 
that of Williston, will in time have to be considered seriously. 

In conclusion, my thanks are due to the officers of the British 
Museum (Natural History), in particular to Dr. W. E. Swinton, for 
affording me the facilities I have enjoyed in my study of the material 
in their charge. Professor D. M. 8. Watson’s critical interest has 
been very encouraging. To the University of Stellenbosch I am 
indebted for a small grant, which has enabled me to come to London. 
My wife has been responsible for the execution of the greater number 
of the figures which illustrate this paper. 


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ANNALS 


AFRICAN MUSEUM 


VOLUME XXXI. 


5 Pac ocecln a New Genus of Cystids from the Lower 
evonian of South Africa. By Joan V. L. Rennie, M.A., 

)., Department of Geology, Rhodes University College, 
IT unstown, South Africa. aie Plate XXXVI.) 


CEL 


ch : PRINTED | oe THE 
ES OF THE SOUTH AFRICAN MUSEUM 


_, AND THE 


( 269 ) 


8. On Placocystella, a New Genus of Cystids from the Lower Devonian 
of South Africa.—By Joun V. L. Renntz, M.A., Ph.D., Depart- 
ment of Geology, Rhodes University College, Grahamstown, 
South Africa. 

(With Plate XXXVI.) 


The class Cystoidea was recorded for the first time from South 
Africa in 1925, when Dr. F. R. C. Reed described a single specimen 
from the Bokkeveld Series (Lower Devonian), collected by him at a 
roadside cutting near Bufielskraal, between De Doorns and Tunnel 
Siding in the Hex River Pass. The specimen was named Placocystis 
africanus sp. nov. (Reed, 1925, p. 30, pl. iv, fig. 1) and was compared 
with the Silurian (Wenlock) species Placocystis forbescanus de Koninck 
(see Bather, 1900, part iii, p. 51, fig. xiii) and other representatives 
of the Anomalocystidae, a primitive but specialised family of echino- 
derms rare in post-Silurian deposits. In 1928 Dr. L. D. Boonstra 
and the writer collected two specimens of a cystid in the First or 
Basal Shales of the Bokkeveld Series at the northern entrance of 
Gamka Poort in the division of Prince Albert. The writer was able 
to make an examination of Reed’s specimen in the Sedgwick Museum 
in Cambridge, and to compare directly with it the two specimens 
here described. 

The writer is indebted to the Research Grant Board of the Union 
of South Africa for financial assistance and wishes to express his 
thanks to Dr. John Hewitt, Director of the Albany Museum in 
Grahamstown, for the use of camera and dark room. 


Placocystella capensis gen. et sp. nov. 


The two specimens from Gamka Poort, here described as Placo- 
cystella capensis gen. et sp. nov,* are preserved as moulds, coated 
with reddish-yellow ferruginous matter, in a greenish micaceous 
mudstone. In each case what is presumed to be the convex external 


* This paper was read at a meeting of the Royal Society of South Africa in 1932, 
and the generic name appeared in the title of the paper published in the minutes 
of proceedings in 1934 (Trans. Roy. Soc. 8. Afr., xxii, p. lvi). The name Placo- 
cystella Rennie 1934 is a nom. nud. 

VOl, XXXI, PART 3. 24 


270 Annals of the South African Museum. 


mould of the concave or right side of the theca is shown, the edge of 
the convex surface on the hand specimens being reflexed upwards 
if preserved at all; in the paratype (S.A.M. 9700) the adcolumnal 
margin is also reflexed upwards, but perhaps as the result of fracture; 
in the holotype (S.A.M. 9701) the lower margins of the two ad- 
columnal plates curve downwards, this feature apparently being an 
original one, and supporting the interpretation of the specimens as 
external moulds. The preservation is relatively superior, though 
the column and brachioles are not attached in either case. There 
is some indication, shown by irregularities of the surface, of shght 
relative movement of plates subsequent to the death of the animal, 
but on the whole there has been little compression of the thecae, 
the majority of the plates being in position and the sutures being 
very clearly marked. A number of the sutures are preserved as 
strong raised ridges, such as could have been formed by infilling; 
the fact that they are raised would also indicate that the specimens 
are external moulds. 

The Holotype of Placocystella (Plate XXXVI, figs. 4-6).—The 
arrangement of the plates is more distinct in one of the above 
(S.A.M. 9701) than in the other, and the specimen is accordingly 
taken as holotype. The moulds of the plates are arranged on a gently 
convex surface. The periphery is intact except along the adoral 
margin. The theca is subquadrate in outline, somewhat wider 
towards the columnal end than towards the oral end, the widest part 
situated ata little less than one-third of the length of the theca from 
the columnal end. The theca is distinctly longer than wide. The 
lateral margins are gently convex, but straightened towards the oral 
end. The adoral margin is not preserved. The adcolumnal margin 
is concave for the attachment of the stem. The margin of the lower 
left lateral marginal plate is abruptly reflexed upwards, the other 
lateral marginal plates not showing this feature. 

The concave side of the theca appears to have been composed of 
twelve plates, ten of them marginals, with two somatic plates en- 
closed by them, the twelve arranged symmetrically about a line 
drawn from front to back. At the columnal end are two large 
marginals, meeting along the middle line, truncate above against 
the larger somatic plate, in length somewhat less than one-third of 
the length of the theca, the sutures between them and the adjacent 
marginals reaching the lower angles of the theca; towards the lower 
angles of the theca these plates are truncate, while the columnal 
margin is fairly deeply embayed for the attachment of the stem, 


On Placocystella, a New Genus of Cystids. 271 


with the edges of the plates curved downwards within the embayment. 
On each side are three marginals, subequal, the lower pair sub- 
triangular, the middle and upper pairs subquadrate. At the oral 
end two smaller marginals meet along the middle line; the one on 
the right side meets the uppermost right lateral in a suture which is 
apparently directed towards the right upper angle of the theca, the 
corresponding suture on the left side being only visible in part. 
Centrally placed is a large subquadrate somatic plate, bounded below 
by the adcolumnal marginals and laterally by the two lower pairs 
of lateral marginals. The second somatic plate is considerably 
smaller than the other, is subquadrate, and placed along the middle 
line. 

The sutures between the plates, as distinguished above, are for the 
most part visible as straight ridges or lines of whose nature there 
can be no doubt. This applies particularly to the sutures between 
the marginals. The sutures bounding the somatic plates are some- 
what disturbed, owing probably to a slight relative movement of 
the plates, but their positions can be determined with some certainty. 

A few of the plates are traversed by ridges, which are not regarded 
as sutures, since (i) they bear little or no relation to the symmetry 
of the theca, (11) they are in some cases much less evident than the 
sutures, (iii) they often pursue an irregular course across the plates, 
and (iv) similar features in the paratype are differently placed. 
Hach adcolumnal marginal is traversed by a faint ridge in the 
anterior-posterior direction, but the position of that on the right is 
quite different from that on the left, and that on the right is curved 
in an irregular manner. A similar ridge pursues a somewhat irregular 
course across the lower corner of the right adcolumnal plate and is 
continued across the lowermost lateral marginal roughly parallel to 
its outer margin, almost certainly a fracture from its appearance 
under the binocular microscope; a ridge is present in nearly the 
same position on the corresponding marginals of the left side. The 
smaller somatic plate is traversed by two faint ridges, one curved 
and almost along the middle line, the other across the upper left- 
hand corner. The remaining plates are free of these features. The 
larger somatic plate has a broad raised band placed obliquely 
across it. 

The three marginals on either side are traversed by irregular, 
relatively broad, wavy, or discontinuous bands placed obliquely to 
the outer margin of the theca. These may have been original 
features of the theca. 


272 Annals of the South African Museum. 


The stem is absent, as well as the brachioles. The margin of the 
lowermost lateral marginal on the left side is curved upwards, 2.e. 
there is preserved a fragment of that part of the plate which bounds 
the rim of the theca; the curved fragment is demarcated by a groove 
(presumably secondary) and traversed by wavy bands. 

Dimensions: length 165 mm.; greatest width 13-5 mm.; length 
of adoral margin about 10 mm. 

The Paratype of Placocystella (Plate XXXVI, figs. 7—-9).—The 
second specimen from the same locality (S.A.M. 9700) agrees with 
the first in dimensions and structure and is taken as paratype of the 
species. The moulds of the plates are arranged on a gently convex 
surface and the periphery is well preserved. The outline agrees 
completely with that of the holotype, with the addition that the 
adoral margin is very slightly convex and the upper corners of the 
theca are rounded angles of about 115°. 

The sutures are for the most part well marked. The sutures 
between the marginal plates are clearly marked on both sides and 
correspond exactly with those of the holotype. At the oral end 
there are only two marginals, that on the left being clearly demarcated 
by a suture along the middle line and a suture joining the periphery 
just behind the left upper corner of the theca, the corresponding 
suture on the right being less defined. The suture between the 
large central somatic plate and the adcolumnal marginals is only 
partly seen, and that between the adcolumnals is not well marked. 
The suture between the two somatic plates is very evident. The 
theca is in various places traversed by faint incisions or low well- 
defined ridges, or broader bands, all of which are probably secondary 
features; noteworthy is the fact that none of these features corre- 
spond exactly in position with similar markings on the plates of the 
holotype. 

The margins of the lateral marginal plates are curved upwards, 
i.e. there is preserved the cast of the rim of the theca on either side. 
The lateral marginal plates are traversed by wavy bands roughly at. 
right angles to the periphery of the theca. In places these appear 
as short crescentic lines, sometimes continuous with one another. 
The stem is absent; the adcolumnal margin is reflexed upwards, 
but apparently as the result of fracture. The brachioles are not 
present in position, but there is a cast of a cylindrical spine-like body, 
8 mm. long, emerging from beneath the theca on the right side. * 

Dimensions: length 17 mm.; greatest width 13-5 mm.; length of 
adoral margin 9 mm. 


On Placocystella, a New Genus of Cystids. 273 


Affinities.—The specimens described above are similar to but 
somewhat larger than the type of Placocystis africanus Reed (1925, 
p. 30, pl. iv, fig. 1) from the same formation, from a locality about 
100 miles distant. In the latter the column is present in position, 
and what is apparently a fragment of a brachiole is preserved near 
the right anterior corner. The theca has much the same proportions 
as Placocystella capensis, but the arrangement of the plates, as in- 
terpreted by Reed, is entirely different from that of the corresponding 
surface of either the new species here described or of Placocystis 
forbesianus de Koninck. Moreover, Placocystella capensis differs 
from de Koninck’s genotype in features of sufficient importance to 
warrant the erection of a new genus. 

The holotype of Placocystis africanus (see Plate XXXVI, fig. 1) is 
too poorly preserved to allow of a definite statement as to the re- 
lationship of that species with the new species here described. The 
figure published by Reed is a drawing of the test apart from the 
matrix on which it is situated, enlarged x24, giving the impression 
of an extraordinarily well-preserved specimen; the sutures in the 
drawing are very well marked and give the author’s interpretation 
of the test, which is described in some detail. The writer examined 
this specimen, which is in the Sedgwick Museum,* and came to the 
conclusion that only a few of the markings present could be inter- 
preted as sutures. The specimen is preserved on a piece of mudstone, 
and has a weathered flaky ferruginous surface. Presumably, from 
the few sutures present as well as from a comparison with the Gamka 
Poort specimens, the concave side of the theca is shown, but in a 
flattened condition. According to Reed this side is made up of some 
twenty plates arranged for the most part in five transverse series in 
four vertical rows. The writer came to the conclusion that the 
arrangement of the plates is to a certain extent very doubtful; that 
few of the markings accepted as sutures by Reed are either as clearly 
marked or as regular as his figure indicates; that several of these 
markings could equally well have been accepted as cracks through 
the plates or the matrix, or lines due to the partial removal of surface 
flakes by weathering. Indeed, the whole surface of the thecal portion 
of the specimen is in a most unsatisfactory condition, rough with 
ferruginous flakes, and marked by a multitude of relatively coarse 
and finer lines of a most irregular kind, clearly related to the weather- 
ing of the surface. There are, however, several relatively straight 
lines on the surface, which are quite unmistakably sutures, and it is 

* Recently numbered A. 3044. 


274 Annals of the South African Museum. 


noteworthy that these correspond in position with the sutures 
observed in the Gamka Poort specimens. Sutures bounding three 
marginals on the right side are distinct, the plates having the same 
shapes and relative positions as in the new species. There is a well- 
marked suture between the right adcolumnal plate and the somatic 
plate anterior to it. The lowermost of the three right marginals is 
traversed by a crack which continues into the matrix, accepted as 
an additional suture by Reed. Indications of portions of a few other 
sutures are present. The writer was unable to find evidence sug- 
gesting a fundamental difference in structure between the two 
Bokkeveld species, but as far as the evidence goes, it would appear 
not more than probable that they belong to the same genus and 
possibly to the same species. In view of the very great imperfection 
of the holotype of Placocystis africanus, and the absence of any satis- 
factory evidence as to the arrangement of the plates in the middle 
and anterior part of the concave side of the test, 2.e. over about two- 
thirds of the exposed surface, the writer has thought fit to propose a 
new specific name for the specimens from Gamka Poort. 

Placocystis africanus must certainly be removed from the genus 
Placocystis on either interpretation of the specimen. Placocystella 
capensis is distinguished from Placocystis forbesianus chiefly by reason 
of its almost perfect bilateral symmetry. In P. forbesianus (see 
Plate XXXVI, fig. 2) there are thirteen plates on the concave side, 
including three adoral marginals; the larger somatic plate is situated 
on the anterior half of the theca, and the smaller somatic plate 1s 
placed towards the left upper corner. The perfect bilateral symmetry 
in the arrangement of the plates thus distinguishes Placocystella from 
the Silurian Placocystis as well as from the early Devonian Ateleo- 
cystis Billings and Anomalocystis Hall (for references see Reed, 1925, 
p. 31). According to Bather (1900, p. 49) the evidence suggests that 
the evolution of the family was towards greater bilateral symmetry, 
and therefore started from the usual sac-like form. Placocystella 
is thus both the most advanced and one of the last representatives 
of the Anomalocystidae. 


On Placocystella, a New Genus of Cystids. 275 


REFERENCES. 


BatuHer, F. A. (1900). Anomalocystidae, in Lankester’s Treatise on Zoology, 
pt. ii, pp. 49-51. 

Hatt, J. (1859). Anomalocystites, in Palaeontology of New York, vol. iii, pp. 132, 
133, 145, 146. 

Kirk, E. (1912). “The Structure and Relationships of certain Eleutherozoic 
Pelmatozoa,”’ Proc. U.S. Nat. Museum, vol. xli, pp. 1-137, pls. i-xi (Anomalo- 
cystidae, pp. 21-29). 

REED, F. R. C. (1925). “Revision of the Fauna of the Bokkeveld Beds,” Ann. of 
the S. African Museum, vol. xxii, pp. 27-225, pls. iv-xi (Placocystis africanus, 
p. 30, pl. iv, fig. 1). 

ScHucHERT, C. (1905). “‘On Silurian and Devonic Cystidae and Camarocrinus,”’ 
Smithsonian Misc. Coll., vol. xlvii, pp. 201-272, pls. xxxiv—xliv (Anomalo- 
cystidae, pp. 204-208). 

ScHUCHERT, C. (1913). Anomalocystidae, in Maryland Geological Survey, Lower 
Devonian, pp. 228, 229. 

Woopwaprp, H. (1880). ‘‘ Notes on the Anomalocystidae, etc.,’’ Geol. Mag., N.S., 
dec. 2, vol. vii, pp. 193-201, pl. vi. 


EXPLANATION OF PLATE XXXVI. 


Placocystis africanus Reed. 


1. Holotype, x 2:5 approx., in the collection of the Sedgwick Museum (A. 3044), 
re-drawn by Miss E. T. Talbot under the writer’s direction. 


Placocystis forbesianus de Koninck. 
2. Concave side of theca, after Bather, modified. 


Placocystella capensis gen. et sp. nov. 


3. Concave side of theca, showing probable arrangement of plates. 
4. Drawing of holotype, natural size. 

5. Holotype, 8.A.M. 9701, x 2-6. 

6. Another photo of holotype, x 2-9. 

7. Drawing of paratype, natural size. 

8. Paratype, S.A.M. 9700, x 2-6. 

9. Another photo of paratype, x 2-5. 


: Pins. ‘ 7 
yy i 


Ps... 


i gts Sy) aa 


ig a A a Date 


Ann. S. Afr. Mus., Vol. XXXI. Plate DVO. ANS | 


L. T. Talbot, del., et J. V. L. R., phot. Neill & Co., Lid. 
CYSTIDS FROM DEVONIAN OF SOUTH AFRICA. 


@ 2) 


9. Lower Cretaceous Lamellibranchia from Northern Zululand.—By 
Joun V. L. Renniz, M.A., Ph.D., Department of Geology, 
Rhodes University College, Grahamstown, South Africa. 
Together with an Account of the Geology of the Cretaceous Beds 
and a Preliminary Analysis of the Associated Ammonite Fauna, by 
S. H. Haveuton, B.A., D.Sc., Director of the Geological Survey 
of the Union of South Africa. 


(With Plates XX XVII-LV.) 


CONTENTS. 
PAGE PAGE 
1. INTRODUCTION : 5 . 277 | 3. PRELIMINARY ANALYSIS OF 
Historical . d 5 te AMMONITE Fauna (by 


SRE ene) on ery 


New Collections from N. 4. Tor LAMELLIBRANCH Fauna . 297 


SETS os : ee Note on Certain Uitenhage 
Acknowledgments and Pre- Species . : 5 . 303 
liminary Remarks . . 282 | 5. DESCRIPTION OF THE SPECIES. 304 
6. REFERENCES . ; : . 388 

2. GEOLOGY OF THE CRETACEOUS 7. TABLE SHOWING DISTRIBUTION 
Beps (by 8S. H. H.) . . 283 OF SPECIES . : at end 


1. INTRODUCTION. 


Historical.—In South Africa marine deposits of Cretaceous age 
are well represented, and have provided the material for a formidable 
list of palaeontological publications. The deposits are all but con- 
fined to the south-eastern and eastern coastal belt, and occur inter- 
mittently for a distance of over 700 miles in the Union and extend 
northwards into Portuguese Hast Africa. General accounts of these 
deposits have been given in recent years by du Toit (1926, pp. 292- 
330), Rogers and Haughton (1929, pp. 143-148), and of the occur- 
rences in Portuguese territory by de Andrade (1929, pp. 147-165). 

In the southern part of the Cape Province only the Neocomian 
is represented, in the Sundays River Beds of the Uitenhage Series, 
which have yielded a rich fauna, monographed by Kitchin (1908), 
and including ammonites which have enabled Spath (1930) to deter- _ 


278 Annals of the South African Museum. 


mine the age more precisely as Upper Valanginian. The Neocomian 
stage has not hitherto been recorded from between the neighbourhood 
of Algoa Bay and Mogambique, a distance of well over 1600 miles. 
A few Neocomian species have been reported from the northern part 
of Portuguese Hast Africa near Mocambique (de Andrade, 1929, 
p. 158), and the well-known Tendaguru Formation in Tanganyika 
has yielded a fine Lower Cretaceous fauna, described by Lange (1914), 
Dietrich (1933), and other workers. 

Between Algoa Bay and Hast Pondoland, for a distance of about 
280 miles, the Cretaceous System is practically absent, and the 
Karroo and Cape Systems outcrop at the coast. In Hast Pondoland 
the Umzamba Beds, almost confined to beach outcrops over some 
25 miles of coast, including outcrops at Umpenyati in southern 
Natal, are characterised by an exceptionally rich fauna of Upper 
Campanian age, described by Woods (1906), Spath (1921 a, 1922), 
Rennie (1930), and others. The beds here rest directly on the pre- 
Karroo basement. The same horizon occurs as sub-outcrops farther 
north at Durban. 

Cretaceous beds reappear at the coast near Port Durnford, south 
of Richards Bay and about 80 miles north of Durban, and stretch 
continuously northwards through Zululand for about 150 miles, 
passing into Portuguese territory, where the system is widespread. 
A considerable portion of the Cretaceous succession is represented 
in the Cretaceous of Zululand, though much remains to be done 
before the sequence is thoroughly understood. The beds rest un- 
conformably on the Karroo System and on Old Granite, for a long 
distance directly overlying the Lebombo Volcanics, and are practically 
undisturbed, the beds dipping eastwards towards the coast at very 
small angles. Fossiliferous outcrops occur at certain favourable 
localities, the greater part of the area covered by the system being 
hidden beneath recent deposits. 

Nearly all the species described up to the present from the Zulu- 
land Cretaceous come from two areas, south of the Umfolosi River 
and north-west of Lake St. Lucia respectively, situated about 40 
miles apart, the latter some 35 miles south of the big bend of the 
Pongola River. The Lamellibranchia described below come from 
localities north of the big bend of the Pongola River, and between 
that river and the Swaziland border. 

A rich fauna has been described by Etheridge (1904) and Spath 
(1921, pp. 224-272), with a few additions by Newton (1909) and 
Rennie (1930, pp. 236-250), from a limited area just south of the 


Lower Cretaceous Lamellibranchia from Northern Zululand. 279 


Umfolosi River, including the well-known fossiliferous locality of 
Umkwelane Hill and the neighbouring localities Lake Itesa (Eteza, 
Isitesa), and “Railway Cutting between Umfolosi Station and Mill 
Halt.” The work of Spath (ibid., pp. 264-272) has made it clear 
that the great majority of, if not all, the ammonites are Campanian 
(and Maestrichtian?), though it is hardly safe to assume the complete 
absence of Lower Senonian horizons; there is no indication of pre- 
Senonian species. The beds here are approximately equivalent to the 
Umzamba Beds in Pondoland, and the transgression of the Senonian 
over older horizons is evident, for at Umkwelane Hill the Senonian 
rests directly on Lebombo basalts. 

A considerable succession of Cretaceous beds is known from the 
area north-west of Lake St. Lucia, where fossiliferous outcrops occur 
along the Umsinene River and its tributary the Manuan Creek 
(Munyuana), which flow into the north-west corner of False Bay, 
Lake St. Lucia. Marine fossils have been described by Etheridge 
(1907), Crick (1907), Newton (1909), Spath (1921, pp. 273-309), 
Rennie (1930, pp. 236-250), van Hoepen (1929, 1931), and Heinz 
(1930). According to Spath the ammonite evidence proves the 
presence of the Middle and Upper Albian, the Cenomanian, the 
Lower Senonian (Coniacian), and the Upper Senonian (Campanian, 
including Maestrichtian?) horizons. An account of the succession 
along the Umsinene River, from above its junction with the Manuan 
Creek to the shore of False Bay, has been given by van Hoepen 
(1926), who divided the sequence into seven beds, to which names 
were given as follows, with suggested correlations :— 


7. Umzamba Beds . Upper Senonian. 
6. Itweba Beds ; . Middle Senonian. 
5. Peronceras Beds ; . Lower Senonian. 
4. Munyuana Beds . Turonian. 

3. Skoenberg Beds ; . Cenomanian. 

2. Umsinene Beds . 

1. Ndabana Beds | Albian 


The palaeontological evidence on which these correlations were made 
has only been published in part by van Hoepen (1929, 1931). The 
presence of the Turonian has not been recognised by previous workers 
(Spath, 1921, p. 268; du Toit, 1926, pp. 316, 319), its presumed 
absence being in conformity with evidence from Angola and else- 
where; the transgression of the Senonian on to pre-Cretaceous forma- 
tions at Umkwelane Hill and in Pondoland points to a break within 


280 Annals of the South African Museum. 


the Upper Cretaceous of Zululand which has not yet been disproved. 
However, van Hoepen (1929 a) insists that there is a practically 
unbroken sequence from the Aptian to the Senonian, and Heinz 
(1930) has identified species of Inoceramus from the Umsinene River 
as Turonian. In a later publication van Hoepen (1929 a, p. 10) 
correlates the Ndabana Beds as Aptian, though without giving 
palaeontological evidence. 

A few Cretaceous species have been recorded from localities in 
Northern Zululand, between the Manuan Creek and the Portuguese 
border, but practically nothing was known of the succession in this 
stretch of country until the visits in 1933 of Mr. H. F. Frommurze 
and Drs. 8S. H. Haughton and A. W. Rogers, whose collections have 
afforded material for this paper. Spath (1921, p. 221) has recorded 
Albian and Senonian ammonites from the Mkusi (Mkuzi) River, and 
Heinz (1930, p. 685) has recorded from the same area a species of 
Inoceramus, said to be Upper Turonian; this locality, 20 miles north 
of the Umsinene, forms an interesting link between the Umsinene 
and Pongola areas. The writer (Rennie, 1930, p. 239) has placed 
on record the occurrence of the Albian species Trigonia cricki, Neithea 
quadricostata, and Veniella etheridge: at the Ingwavuma road drift on 
the Pongola River, as well as Nordenskjéldia natalensis and Pleuromya 
africana, Senonian species, at some locality west of the Pongola River. 

The Cretaceous of Zululand is continuous with that of the district 
of Lourengo Marques in Portuguese Hast Africa, where outcrops occur 
from the Zululand border northwards as far as the neighbourhood 
of the point of entry of the Limpopo River into Portuguese territory, 
a distance of about 300 miles. Outcrops occur on the Maputo (Usutu) 
River near Catuane, close to the Zululand border, and between Bela 
Vista and the east coast to the south of Delagoa Bay; Cretaceous 
beds are known to extend northwards along the eastern flank of the 
Lebombo Mountains as far as the Limpopo, where they rest on the 
Lebombo Volcanics and dip eastwards at small angles beneath recent 
deposits as they doin Zululand. The area is at present being investi- 
gated by the officers of the Department of Mines and Industries 
of Mogambique. It would appear that at least the Aptian, Albian, 
Cenomanian, and Senonian stages are present. A number of species 
of Aptian ammonites have been recorded by Kilian (1902), Krenkel 
(1910), and Spath (1921, pp. 309-318) from Catuane, Powell’s Camp, 
and other localities (de Andrade, 1929, p. 159) in the region south 
of Delagoa Bay; the Aptian has not hitherto been recorded from 
farther south in Zululand, unless van Hoepen (1929 a, p. 10) is 


Lower Cretaceous Lamellibranchia from Northern Zululand. 281 


correct in correlating his Ndabana Beds at the base of the sequence 
on the Umsinene with that stage; according to Spath the Aptian 
beds at Catuane, etc., are probably Upper Aptian. From Catuane 
Spath (1925, pp. 180-196) has also described a number of Upper 
Albian ammonites, while Cox (1925, pp. 202-207) has described some 
gastropods from the same locality and horizon. The Senonian stage 
has been recognised at Incomanini on the Incomati (Komati) River, 
north of Lourengo Marques, the fauna containing a few species of 
Lamellibranchia identical with or allied to species in the Umzamba 
Beds of the Union (du Toit, 1926, p. 327); the Lamellibranchia have 
not been described, but Cox (1925, pp. 202, 208-215) has recorded 
a number of Gastropoda of somewhat Tertiary aspect; ammonites 
appear to be absent. Spath (1925, pp. 196-200) has recorded a 
Cenomanian ammonite and two Upper Senonian ammonites from 
Maputoland, the region south of Delagoa Bay. 

In the region between the Limpopo and the Zambesi the Cretaceous 
System is for the most part covered with recent deposits, but beds 
referable to the system appear here and there, both on the coast 
and inland. Newton (1924) has recorded both Senonian and Danian 
faunas from the Cheringoma (Sheringoma) Plateau adjoining the 
Urema trough, north of Beira; while the Upper Cretaceous species 
Lopha (= Alectryonia) ungulata was recorded by Newton (1896) from 
the neighbourhood of Sofala. 

Still farther north the Cretaceous is absent for a considerable 
distance, but Cretaceous beds reappear on the coast south of Mocam- 
bique, and can be traced into Tanganyika. From this region 
Choffat (1903) has recorded Cretaceous beds at Conducia, apparently 
largely Cenomanian; the Neocomian, Aptian, and Albian stages are 
said to occur in this region also (de Andrade, 1929, pp. 158-160). 

The New Collections from Northern Zululand.—The Lamelli- 
branchia described in this paper come from outcrops in Northern 
Zululand, situated for the most part on the Pongola River and in 
the region between the Pongola River and the borders of the Trans- 
vaal and Swaziland, not far to the south of the southern boundary 
of Portuguese East Africa. Prior to 1933 this tract of country was 
practically unknown from a geological point of view, though it was 
known that the Cretaceous beds exposed north-west of Lake St. 
Lucia extended northwards along the lower course of the Pongola 
and into Portuguese territory. In 1933 Mr. H. F. Frommurze, B.Sc., 
of the Union Geological Survey, while engaged in irrigation work in 
this area, collected from fossiliferous Cretaceous beds along the 


282 Annals of the South African Museum. 


Mfongosi stream, a tributary of the Pongola, and the collections sent 
to Cape Town were deemed of such importance that the area was 
immediately visited by Dr. 8. H. Haughton, now Director of the 
Survey, and Dr. A. W. Rogers, formerly Director of the Survey. 
Cretaceous outcrops were located over a wide area and an attempt 
was made to work out the sequence by careful collections of fossils 
on a zonal basis. Subsequent to their visit further collecting was 
done by Mr. J. 8. Hutt, Surveyor to the Irrigation Department, 
without reference to localities or horizons; this collection was pre- 
sented to the Transvaal Museum. 

Acknowledgments and Preliminary Remarks. — The Lamelli- 
branchia described below form part of the collections obtained by 
Mr. H. F. Frommurze, Dr. 8. H. Haughton, and Dr. A. W. Rogers, 
and by Mr. J. 8. Hutt in 1933. The writer is very greatly indebted 
to Dr. 8. H. Haughton, Director of the Geological Survey of the 
Union of South Africa and Honorary Curator of the Palaeontological 
Collections in the South African Museum in Cape Town, for placing 
the Lamellibranchia at his disposal for description, and for much 
useful advice. Dr. E. L. Gill, Director of the South African Museum, 
and Dr. T. W. Gevers, formerly on the Union Geological Survey, 
have very kindly forwarded to the writer specimens and literature 
contained in the collections in Cape Town, while the writer is parti- 
cularly grateful to Dr. Gill for arranging for the publication of this 
paper and for help in various ways. Mr. L. R. Cox, of the Geological 
Department of the British Museum (Natural History), and Dr. K. H. 
Barnard, Assistant-Director of the South African Museum, have 
given invaluable help by looking up early references to Trigonia not 
accessible in Grahamstown, and giving their opinions on certain 
problems of nomenclature; to them the writer is very greatly in- 
debted. The accompanying plates were prepared by the writer, and 
his best thanks are due to Dr. John Hewitt, Director of the Albany 
Museum in Grahamstown, for granting every facility in the use of 
camera and dark room; and for much helpful criticism. 

The cost of photographic materials was defrayed by a grant from 
the Research Grant Board of the Union of South Africa, for which 
the writer expresses his thanks. . 

The collections obtained by Mr. H. F. Frommurze have been placed 
in the South African Museum in Cape Town; those made by Mr. 
J. S. Hutt in the Transvaal Museum in Pretoria; where individual 
specimens are referred to, “S.A.M.” or “T.M.” precede the catalogue 
numbers. The collections made by Dr. 8. H. Haughton and Dr. 


Lower Cretaceous Lamellibranchia from Northern Zululand. 283 


A. W. Rogers are in the Cape Town office of the Geological Survey of 
the Union, at the South African Museum; these are simply labelled 
with the horizon letter and number, but types and figured specimens 
at least will be transferred to the collections of the South African 
Museum in due course. A few specimens from the Albany Museum 
(A.M.) are also referred to. 

In recording localities in the text the number of available speci- 
mens from each locality has been indicated by a number in brackets, 
e.g. Z19 (4) indicates that 4 specimens were collected at horizon 
Z19. References to published papers have been made by indicating 
the author’s name and the date of publication; a list of references 
will be found at the end of the paper. 

In preparing the descriptions that follow, reference to a number 
of badly preserved, tragmentary, or otherwise doubtful species of 
Lamellibranchia has been omitted. It was considered that no good 
purpose would be served by giving descriptions or figures of these 
species, either because the locality at which they were collected was 
not precisely recorded, or because they were found at one locality 
and horizon only, and therefore fail to be of any value in correlating 
horizons at one locality with those of another, or because of some 
uncertainty as to the identity of the genus to which they belong, 
or because of great imperfection in the preservation of the shells 
concerned. Included in this category are a number of specimens 
referable to the Veneridae, to Ostrea, to Pecten, and to a few other 
genera. | 

Trigomae belonging to the section Scabrae, which would be referred 
to the sub-genus Scabrotrigonia Deecke, or to the several sub-genera 
proposed by van Hoepen (1929), have not been specially dealt with 
here, although several species are represented in the abundant material 
available. Apart from this group, which will be worked out later, 
all the recognisable and significant species of Lamellibranchia con- 
tained in the collections are described below. A brief reference to 
the Scabrae is included, however. 


9. An ACCOUNT OF THE GEOLOGY OF THE CRETACEOUS BEDs. 


(By S. H. Haughton, M.A., D.Sc., Director of the Geological 
Survey of the Union of South Africa.) 


The localities from which the fossils described in the following 
pages were collected lie in the coastal belt of Northern Zululand, in 
the magisterial districts of Ubombo and Ingwavuma. This belt is 
bounded to the west by the Lebombo mountain range, a dissected 


Mzinyeni 
Pan 


NATIVE RESERVE 
NO 15 


~ 
PongoloA 


a 


Mokotini 
A (Camp) 


Lower Cretaceous Lamellibranchia from Northern Zululand. 285 


peneplaned ridge of Karroo volcanic rocks whose structure and 
features have been recently described by du Toit (Trans. R. Soc. S. 
Afr., 1929, vol. xvi, pp. 189-217). The range has an approximately 
north-south strike, and—in the area visited by the writer—its 
wonderfully even sky-line is broken only by the deep gorges of the 
Mkuzi, Pongola, and Ingwavuma rivers. Transecting the range in 
a west-east direction these streams begin to meander as soon as 
they leave the foothills of rhyolite and enter the flat coastal plain, 
the Mkuzi turning in a south-east direction to empty itself into False 
Bay, the Pongola—joined by the Ingwavuma—flowing northwards 
into Delagoa Bay. 

In addition to these major streams which cut through the Lebombo, 
a number of smaller tributaries rise on its eastern flanks. Where 
they, and the bigger streams, flow through the Cretaceous beds they 
form a number of disconnected deep pools in which hippopotami 
and crocodiles are to be found. 

The actual river-beds lie about 60 feet below the general level 
of the coastal plain. This plain is capped by a covering of gravel 
with, in parts, a consolidated calcareous grit of marine origin. Sand, 
too, is common. The fossiliferous Cretaceous beds are only exposed 
on the banks of the streams or sides of gullies. The localities in- 
vestigated are shown on the accompanying plans, and are—from 
south to north—the Myesa Spruit, the Mfongosi Spruit, the Pongola 
River with a tributary gully, the Msinyene Pan, and the Lombag- 
wenya Spruit. Small collections were also made south of Pongola 
Poort at Mokatini Camp and to the south of it. 

The investigation of this area by the writer and Dr. A. W. Rogers, 
F.R.S., was only made possible by the kindness of the Director of 
Irrigation, who kindly placed all his large-scale contour plans at our 
_ disposal, and of Mr. J. 8. Hutt, Surveyor-in-Charge of the Lower 
Pongola Irrigation Survey, whose camp at Otobotini provided us 
with every comfort, and who placed both his time and local knowledge 
enthusiastically at our service. Mr. Hutt, both before and after our 
visit, was active in the collection of fossils, and presented the bulk of 
his material to the Transvaal Museum. Some of it has been studied, 
and the results are incorporated in the palaeontological descriptions 


which follow. 
Description of Localities. 


1. Myesa Spruit.—Very platy rhyolites with a dip of about 10°-12° 
in a direction 10° north of east form a cliff just below the drift on 
VOL. XXXI, PART 3. 25 


286 Annals of the South African Museum. 


the southern branch of the stream. The contact between the 
rhyolites and Cretaceous beds is not seen, being overlain by alluvium 
and gravels. \ | 

North of the drift, across the northern branch, buff sandy beds 
containing gypsum are seen in a roadside hole and in a shallow donga. 
They contain a band of hard gritty quartzite, which is nodular; on 


Rag. We 


the weathered surface this shows masses of fibrous calcite, which 
may possibly be replaced wood. 

At locality M2 are unevenly bedded pebble beds and grits, buff in 
colour, with a dip towards the south-east. The exposures on the 
left bank of Myesa Pan (M3 and M4) are poor. They consist of buff 
soft sandstones with occasional small lenses of hard calcareous grit, 
which sometimes carry badly preserved shells. 

2. Mfongost Spruit.—The most easterly outcrop of rhyolite along 
this spruit is seen at the point where the river leaves the foothills, 
some 3-8 miles in a direction west 30° north from the drift across 
the spruit on the main road. Here the rhyolite is platy and 
amygdaloidal, and dips at 12-15° in a direction slightly north 
of east. 


Lower Cretaceous Lamellibranchia from Northern Zululand. 287 


Below this the river banks are low and gravel-covered. A few 
hundred yards downstream there is a short exposure of consolidated 
boulder beds and gravels which appear to have a slight dip down- 
stream. ‘There is no evidence available as to their age; but none 
of the recent conglomerates along the rivers appear to be so compacted 
as these. 

The most westerly outcrop of Cretaceous beds occurs on the south 
branch of the spruit at a point about 2 miles above the drift, measured 
in a straight line, bearing west 15° north from the drift. This con- 
sists of bufi-coloured, sandy pebble beds and sandy grits in which 
no fossils were seen. 

Just above the junction of the two branches of the spruit there 
is a good exposure on the south branch of a thickness of about 30 
feet of buff sandstones, grits, pebble-washes, and conglomerate, with 
a dip of 3° towards the east. In places current-bedded bands show 
a much higher dip in the same direction. Included in the conglom- 
erate are many rhyolite pebbles, some of agate, and some of sedi- 
mentary rocks. The sandstones carry ferruginised badly preserved 
fossil wood, but no marine fossils were seen. 

At the big cliff, on the left bank of the stream above the drift, there 
is a succession of beds, mostly of buff sandstone with hard lenticular 
nodules about 55 feet thick. This locality was designated Za. The 
basal 5 feet contain six thin hard bands filled with fossils, which 
include Trigonia obesa, Trigonia haughtom, and Trigonia hennigi, 
separated by buff sandy joint clays. Two feet above this is a narrow 
band with the same Trigomia species and a Belemnite. 

The buff sandstones of the higher part of this succession outcrop 
at the first cliff above the drift on the right bank of the river. Here 
some of the gritty sandstone is rather more micaceous than at locality 
Za; the finer-grained sediments contain fragments of stems; and 
a few fossiliferous concretions with Trigonia occur. On the slope 
above the cliff (locality Zo) one specimen of a large uncoiled ammonite, 
Tropaeum cf. gigas, was discovered, and some fragments provisionally 
assigned to the genus Toxoceratoides. 

Below the drift the cliff on the left bank has at its western end 
sandy shales with rounded and irregular nodules containing a species 
of Acanthohoplites and Dicroloma sp. (locality Z1). Then follows a 
succession of sandy clays with thin bands of fossiliferous limestones 
(locality Z2) containing Trigonia obesa, Trigonia pongolensis, Cardium 
rogerst, and Dicroloma sp. The highest horizon in this cliff section 
(Z3) consists of buff argillaceous sandstones with thicker elongate 


(oes ies: 


LII7S COOP OOO? GOO! a Cosa Ss! See. ; Amgen vi 
SHY (PMYE LiMy 
HZ. 
(tee 
oe Se 
a) ——- 
1S) 
= 
: ; 
N f > | 
. Tas . =) ww erty 
ce Au —x~_ OZ 
‘i ye ia TI So AW, 
RY gs Z)_/ Ou sie Ne oN ; 
9 Z i PUB) SOUPS Lf 1 i AS BEE \ 
é TONY \ 
Z; ae 
1S. fees ‘ ae, v, we 


GP ree Re WSs cae S ---<5 


Lower Cretaceous Lamellibranchia from Northern Zululand. 289 


lenticles of limestone, and yielded Trigoniae of the Scabrae group, 
Trigoma obesa, Gervillia dentata, and Cardium rogers. 

On the right bank the first cliff is of buff sandy clays, with plant 
fragments and calcified wood with, at the top, bands of unfossiliferous 
septarian nodules. This cliff is succeeded by a slope, at the top of 
which is a fossiliferous limestone (locality Z4), with Gervillia dentata, 
Pseudavicula (*%) africana, Trigonia obesa, and Trigomae of the Scabrae 
group. Many of the shells here were broken before consolidation; 
further, the limestone carries large mud-pellets and some pebbles. 

This fossiliferous limestone dips at a low angle downstream at the 
top of the next cliff on the right bank. Near the lower end of the 
cliff it gives place gradually to fossiliferous soft sandstones, which 
yield a large number of small shells. The limestone is in reality 
a series of flat lens-like bodies occupying one horizon. Between the 
lenses the grey marly sandstone is fossiliferous. Shells of the same 
species as occur in the limestone are also found some 2 feet below it. 

Locality Z5 occurs on the slopes below a terrace farther down the 
right. bank of the river and shows outcrops of shelly lmestones 
definitely higher in the succession than those of Z4. From here 
were collected Gervillia dentata, Pseudavicula (%) africana, Trigonia 
obesa, Belemnites, and Acanthoplitids. Farther along the slope, 
opposite a cliff of the left bank, dirty bufi-coloured, weathering, shelly 
limestone bands, frequently with spheroidal weathering, become 
common. This is locality Z6. Logs of heavily bored calcified wood 
are numerous, and the fauna contains Gervillia dentata in large 
numbers, Inoceramus concentricus (*%), Trigonia frommurzer, Trigoniae 
of the Scabrae group, Panope gurgitis, Acanthohoplites spp., Toxocara- 
toides sp., and Ancycloceras (Australiceras) sp. 

The higher ground on the right bank of the spruit ends to the 
east in a narrow gravel-capped spur. Locality Z11 is on the south 
flank of this spur; locality Z12 on the north. The outcrops are of 
buff shales and nodular limestone, and the fossils collected include 
a Pseudodiadema, Acanthohoplites spp., several large ammonites, 
Cucullaea woodsi, Pseudavicula (*%) africana, Trigonia obesa, abundant 
Trigonia frommurzer, Trigoniae of the Scabrae group, Protocardia ct. 
sphaeroidea, and Panope gurgitis. 

On the left bank, locality Z7, is a cliff opposite the space between 
Z5 and Z6. A highly fossiliferous limestone band overlies sandy 
clays with limestone concretions. Gervillia dentata, Trigonia from- 
murzei, and large uncoiled ammonites are very common; Isognomon 
sp. and scabrous Trigoniae also occur. 


290 Annals of the South African Museum. 


Above the limestone band which delimits the upper part of this 
succession is a slope formed of soft sandy shales and calcareous shelly 
concretions. This is locality Z8. Numerous bored logs, some of 
them 9 feet in length, occur here. Large loosely coiled Lythceratids, 
Acanthohoplites spp., Lytocrioceras, Cheloniceras, Diadochoceras sp. nov. 
aff. nodosocostatum, Tropaeum cf. bowerbanku, Helicancyloceras (%), 
and Sonneratia all occur in fair numbers, and the Lamellibranch 
fauna includes Trigoma frommurzei, Trigonia obesa, scabrous Trigoniae, 
Cucullaea woodsi, Glycymeris cf. griesbachi, Gervillia dentata, Isog- 
nomon sp. (2), Hxogyra conca, and Panope gurgitis. The band con- 
taining this fauna continues for several hundred yards downstream, 
and is overlain at Z9 by a series of outcrops of ball-like concretions 
that contain very few fossils, succeeded by outcrops of shelly lime- 
stone with minute shells, Trigonia frommurzei, Rhynchonella sp., 
Acanthohoplitids, and Diadochoceras. In these beds a log of calcified 
wood 30 feet long was seen. 

A gully running into the spruit at Z10 exposes soft buff shales 
with numerous logs of wood and a layer of ball-like concretions with 
sparse, large, flat, involute Beudanticeras—like ammonites, Gervillia 
dentata and Trigonia frommurzer. 

Below this gully the cliff shows bands of highly fossiliferous sand- 
stone and sandy limestone dipping at a low angle eastwards. The 
upper end of this cliff (Z13) has Pseudavicula (*%) africana, Trigonia 
frommurzei, Neithea quinquecostata, Protocardia cf. sphaeroidea, 
Douvilleiceras mammillatum, Beudanticeras, and Puzosia cf. stoliczkai; 
the lower end (Z14) yielded Isognomon sp., Inoceramus concentricus (*), 
and Panope gurgitis. 

Locality Z15, which is near the lower end of the Mfongosi Pan, 
yielded a possible new species of Beudanticeras. 

3. Pongola River.—North of the mouth of the Mfongosi Spruit, 
between the Irrigation Survey points RAF and CRAFT, a short 
rather wide valley runs into the Pongola River. Along the sides 
of the valley is a continuous outcrop of a prominent thin band of 
limestone crowded with shells of Turritella manuanensis. This lime- 
stone has an apparent slight dip to the north-east. Below it are 
sandy buff clays with limestone concretions, which are highly fos- 
siliferous, Dipoloceratids, Mortoniceratids, and Desmoceratids being 
common, particularly at an horizon about 40 feet below the Turritella 
limestone. This limestone and the beds below were numbered Z16, 
whilst the beds above the limestone on the valley sides were numbered 
Z17. From Z16 were obtained Puzosia stoliczkar, Hysteroceras spp., 


Lower Cretaceous Lemellibranchia from Northern Zululand. 291 


Dipoloceras quadratum, Deiradoceras spp., Pervinquieria spp., Gervillia 
dentata, Inoceramus concentricus, Scabrotrigonia spp., Pholadomya 
vignest, Gonomya sp., Veniella etheridgei, Exogyra conica, Ostrea sp., 
and Turritella manuanensis. 

The beds Z17 are less fossiliferous, but carry Adkinsites umsinenense, 


SZ Gastergpod 
Liumeslore. 


4000 FEET 


Pervinqueria, Hysteroceras choffati, Dipoloceras cristatum, and 
Phylloceras velledae. 

North of the junction of this valley with the Pongola, on the farm 
Neser, there is a very fine long cliffi-section on the left bank of the 
river. The section is about 40 feet high, and its upper 5 feet is 
formed of Tertiary or Recent gravels and sands. The Cretaceous 
beds consist of a succession of clays and clayey sandstones with 
limestone bands and bands of nodules, and their apparent dip is 
at a low angle downstream. 


292 Annals of the South African Museum. 


Near the base of the section (Z19) is a prominent thin pebble bed. 
Below this the clays are highly fossiliferous, Oxytropidoceras sp. nov., 
Nautiloids, Neithea quadricostata, Cucullea woodsi, Gervillia dentata, 
Trigoma cricki, Protocardia cf. sphaeroidea, Veniella etheridgei, Exogyra 
conica, Turritella manuanensis, and Avellana cf. incrassata being the 
commonest forms present. Above this pebble bed there are definite 
highly fossiliferous layers with the same Lamellibranchs as in the 
lower bed (with the exception of Veniella), together with small 
Dipoloceratids and the echinoderms Hemiaster zululandensis and 
Holaster vanhoepent. 

Towards the top of this section (Z19) Puzosia cf. bhima becomes 
the commonest ammonite, and Dipoloceras cristatum is found. 

Proceeding northwards the cliff section continues (Z20) and some 
25 feet of buff sandy clays and limestone nodules carry abundant 
Exogyra, Vemella, Trigona, Neithea, and Nautiloids. Ammonites, 
however, become rather scarce, Myloceras, Puzosia, and Dipoloceras 
being the only genera collected. 

Still farther to the north the cliff becomes lower, and the post- 
Cretaceous grits and gravels sink considerably in level. The Creta- 
ceous beds are more sandy than to the south, and carry sporadic 
nodules of calcareous quartzite, some of which are fossiliferous. 
Fossils are scarce in the buff sandstones. From this locality (Z21) 
Holaster, Hemiaster, Inoceramus concentricus var. subsulcatus, Phola- 
domya vignesi, and Goniomya are recorded. 

4. Lombagwenya Spruit.—The main road from Otobotini to Ndumu 
crosses this spruit to the west of the Kwambosi Pan, and rises up to 
the plateau past the Lombagwenya Store. On the slopes down to 
the spruit, west and north-west of the store, richly fossiliferous out- 
crops occur; and other exposures of Cretaceous beds occur a short 
distance below the drift. 

Three traverses were made across this fossiliferous slope. 

Immediately to the west of the store a thickness of about 125 feet 
of beds (locality L3) consists mainly of buff sandy marls with cal- 
careous nodules. Near the base are large open-coiled Crioceratids with 
depressed whorl section and prominent widely spaced ribs passing 
right across the venter, associated with Gervillia dentata and Trigonia 
frommurzei. Just above the bed most prolific in ammonites is a 
narrow band of sandy clays, below a thin limestone, that contains 
numbers of Rhynchonella and Terebratula. The upper 50 feet of beds 
carry several species of Douvillecceras including typical D. mamillatum, 
Phylloceras sp., Leymeriella sp.n., Acanthohoplites spp., and a new 


Lower Cretaceous Lamellibranchia from Northern Zululand. 293 


genus of ammonite. At the top of the slope large Terebratulids 
occur. 

Following the slope to the north-west one encounters blocks of 
fossiliferous limestone; here and there are low benches formed of 
limestone outcrops. The buff shaley beds which are intercalated 
with these are only occasionally exposed in shallow dongas. 

The second traverse was made upwards from the southern end 


Iie, Ze 


of a cliff about 2300 yards north-west from Lombagwenya Store. 
The chff is composed of almost unfossiliferous buff sandy clays, 
showing some false-bedding with fossiliferous limestone nodules 
(locality L8). Phylloceras sp. and Hxogyra conica (%) occurred here. 
Just above the crest of the cliff occurred Megacucullaea sp., Trigonia 
hennigi, Tropaeum sp., and higher up the gully are limestones with 
large Ostrea, Panope gurgitis, Aconeceras cf. walshense, and Toxocera- 
toudes of the royert group (locality L9). The next outcrop up the 
sully (L7) is limestones with Pholadomya, Panope, Australiceras ct. 
lampros, Lytoceras cf. mikadiense, Cheloniceras (%), and numerous 
specimens of a sinistral Turrilites cf. emericianus. At the top of the 
slope are specimens of the loosely coiled Crioceratids generally similar 
to those that were found near the base of locality L3. 


294 Annals of the South African Museum. 


The third traverse was made beyond the northern end of the cliff. 
The lowest beds seen here (L10) are buff sandy marls with large 
lenticles and rounded masses of slightly calcareous sandstone carrying 
fossil wood. The lenticles sometimes carry shells, especially a medium 
sized Chlamys. Above these beds is a band of shelly limestone (L11) 
with Panope gurgitis, Aconeceras cf. nisus, Acanthohoplites spp., 
Leymeriella spp. Slhghtly to the south-east, and at about the same 
height above the river, are shelly sandstones (L12) with Cucullaea 
woods (%), Megacucullaea sp., Trigonia hennigi, but no ammonites. 
Above these is a band of sandy shale with fragile Scabrotrigonia, 
Gervillia dentata, Ostrea sp., and Thetironia sp., followed by shales 
with limestone and very abundant fossil wood. Higher is a zone 
of buff shales with limestone concretions yielding enormous specimens 
of Gervillia, Trigonia hennigi, Ostrea sp., Nautilus, and Phylloceras, 
with fossil wood (L4). Above this is an outcrop of shelly limestone 
with large loosely coiled Crioceratids, whilst at the top of the slope 
are numerous weathered, rounded clay nodules (L3+) * of various 
sizes, many of which are fossiliferous and carry Isognomon sp., 
Lopha diluwana, Tropaeum cf. mozambiquense, Tropaeum cf. arcticum, 
Australiceras, Acanthohoplites of the roseanus group. 

Exposures of Cretaceous beds occur also below the drift. 

On the right bank of the spruit about 150 yards below the drift 
is an exposure of a thickness of about 30 feet of stiff sandy clays with 
one layer of hard fossiliferous limestones and some sporadic calcareous 
nodules (locality L6). These yield Phylloceras, Douvilleiceras, Scabro- 
trigonia, Terebratula, Exogyra cf. conica, and Gervillia dentata. 

On the north side of the spruit are low ridges above the alluvial — 
plain. On the slopes of these (L1) are occasional outcrops of weathered 
calcareous nodules that contain large specimens of Gervillia dentata, 
Scabrotrigonia, Panope gurgitis, Glycimeris, brachiopods, Douvilleiceras, 
and Oxytropidoceras (*) sp. juv. 

Farther down the spruit a low cliff above the water has buff clays 
which are highly fossiliferous, but retain the fossils only in the form 
of internal casts. Fragments of ammonites only were seen here 
(L.2), one small crushed Echinoid, and Scabrobrigonia sp. 

5. Msinyene Pan (locality L5).—The low slopes on the western 
side of the south end of this pan show few outcrops; but some fossils 
were collected here which are of interest. Among them was a speci- 
men of a new ammonite belonging to the form already recorded 


* Locality L3t is distinct from L3 and may not be the same horizon. L37 is 
equivalent to [L3] in fig. 4. 


Lower Cretaceous Lamellibranchia from Northern Zululand. 295 


from the upper beds of section L3, Leymeriella sp., and a species 
of Beudanticeras similar to one found in the highest beds of the 
Mfongosi Spruit. Scabrotrigonia sp. also occurs, and an imperfect 
shell which Rennie has tentatively assigned to Neithea quadricostata. 

6. Mokatini.—The area to which this name has been given lies 
between the Pongola and Mkuzi Rivers. Apart from the rhyolite 
foothills of the Lebombo the country is almost entirely covered 
by soil and sand, and shows very few outcrops of the Cretaceous beds. 

The low ridge which runs south from the Mokatini Irrigation 
Survey camp, on the south side of the Enseleni road, carries an 
exposure of a band of very hard shelly limestone from which it is 
difficult to extract shells (locality Z24). The shells are arranged in 
layers separated by bands of hard, barren, brown-weathering, fine- 
grained calcareous quartzite. From here come Pseudavicula (?) 
africana, Scabrotrigoma sp., and Thetironia sp. 

About 4 miles to the east of this, and at an altitude about 150 
feet lower, is a small exposure of a hard limestone with pebbles and 
shell fragments and blocks of fossil wood. A little downstream from 
this.a softer shaley bed (Z25) yielded Trigonarca cf. ligeriensis, 
Exogyra conica, Trigoma blanckenhorni, and Veniella etheridge. 


3. A PRELIMINARY ANALYSIS OF THE AMMONITE FAUNA. 


(By 8. H. Haughton, M.A., D.Se., Director of the Geological 
Survey of the Union of South Africa.) 


Until the ammonite fauna is identified specifically exact determina- 
tion and correlation of the various horizons present will not be 
possible. Nevertheless, certain generalisations may be made which 
are of interest. 

1. Mfongosi Spruit Succession.—It is estimated that a thickness 
of 800 feet of fossiliferous beds are exposed along this spruit, the 
lowest being exposed at locality Za and the highest at locality Z15. 
On the right bank ammonites were collected at, in ascending order, Zo, 
25, 78, and Z12: on the left bank at Z1, Z7, Z8, Z9, Z10, Z13, Z15. 

The lowest horizon (that at Zo) yielded Tropaeum cf. gigas, and 
must lie some 30 feet below Z1, which carries Acanthohoplites sp. nov. 
The presence of Tropaewm would seem to indicate the presence 
of the top of the Lower Aptian or of the lower part of the martini 
zone of the Upper Aptian at Zo. 

The assemblage from Z8 includes forms that may be representative 
of more than one of the European zones. Tropaeum ct. bowerbanki 


296 Annals of the South African Museum. 


is compared with a species that 1s representative of the middle sub- 
zone of the Tropaeuman of the Upper Aptian. On the other hand, 
Diadochoceras sp. nov. aff. nodosocostatum (ident. L. F. Spath) is 
compared with a species that is characteristic of the lowest zone 
of the Albian. It is possible, however, that this apparent mingling 
may be susceptible of interpretation, as the locality Z8 extends for 
nearly 1500 feet along the side of the spruit and contains a thick- 
ness of some 78 feet of beds. Further collecting may show that 
each of the critical forms is confined to a definite horizon, and that 
the intervening subnodosocostatum zone may also be present. It 
is of interest that a species of Acanthohoplites, comparable with 
A. aschiltaensis, comes from the lower beds at Z9; this species is 
characteristic of the upper part of the swhnodosocostatum zone. 

At locality Z9 Acanthohoplites bigourets is definitely indicative of 
the presence of the lowermost Albian (Clansayes horizon); and at 
Z13 Douvilleiceras mammillatum shows the presence of the base of 
the Middle Albian (mammullatus zone). Here again there is a hiatus 
in the faunal succession as worked out in Europe, corresponding to 
the Leymeriellan zone of the Lower Albian, which may be represented 
by the beds at Z10, which contain Beudanticeras sp. These beds 
cannot be more than a few feet thick. 

Since no stratigraphic break was visible in the rock succession, 
it may be accepted that along this section there is a succession of 
fossiliferous beds ranging from the lower part of the martini zone 
of the Upper Aptian at least to the mammullatus zone of the Middle 
Albian, and that the unfossiliferous beds in the upper reaches of the 
spruit are Lower Aptian and may range down into the Neocomian. 
A tentative classification of the beds, according to the locality numbers, 
is as follows :— 


Middle Albian . mammillatus zone INS; LAD: 
tardefurcata zone Z10 (2), Upper beds 
(Leymeriellan) of Z9 (2). 
ee nodosocostatum zone Z9. Upper part of 
(Acanthohoplitan) Z8 (2), Zz 
subnodosocostatum zone Lower part of Z9? 
(Parahoplitan) Lower part of Z8? 
Upper Aptian . i (2);) ZS eee 
ZT (2). 
martini zone Zo, Ll (2). 


(Tropaeuman) 


Lower Cretaceous Lamellibranchia from Northern Zululand. 297 


2. Lombagwenya Spruit.—Beds of approximately similar age to 
the above occur in this area. The presence of the mammillatum 
zone is evidenced by abundant Douvilleiceras in the upper 50 feet 
of the L3 succession; these upper beds also yielded Leymeriella, so 
that it may be possible to see in them lower tardefurcata zone beds 
and upper mammillatum zone beds. The lowest beds in this section 
contain Crioceratids that are comparable with those of Z7 and ZS. 

The assemblage at L11 contains peculiar features, in that Aconeceras 
and Leymeriella apparently occur together. Hitherto Aconeceras has 
been considered to be confined to the Gargasian, with the exception 
of possible representatives from the Bedoulian, but is unrecorded 
from the Albian; Leymeriella, on the other hand, seems to be con- 
fined to the upper half of the Lower Albian. It is certain that L11 
lies below L3y7, in which Gargasian Ancycloceratids and Acanthohoplites 
of the roseanus group occur. It may be remarked that the associa- 
tion of Tropaeum with Australiceras both at L3} and at Z6-Z8 tends 
to throw doubt on Whitehouse’s suggestion that the former was a 
descendant of the latter. 

3. Pongola River.—In this area the fauna is definitely Middle and 
Upper Albian. The lowest horizon (basal part of Z19) contains 
Oxytropidoceras, whilst Manuaniceras occurs just above the pebble 
bed near the base of this section. Slightly higher were found 
Dipoloceras cristatum, Dipoloceras bouchardianum, and Puzosia cf. 
bhima. The Dipoloceratids of this assemblage are closely compar- 
able with those from the top of the Middle Albian of England, the 
cristatum sub-zone of the Lower Gault. 

Localities Z16 and Z17 contain forms that are indicative of the 
Upper Albian, such as various species of Pervinqueria and Deirado- 
ceras. The former ranges in Hngland from the orbigny: to the 
aequatorialis sub-zones. Hysteroceras, which is also fairly abundant 
here, is typical of the lower part of the Upper Albian. On the other 
hand, Dipoloceras cristatum has also been found at this locality, as 
has Adkinsites umsinenense. The presence of these probable members 
of the cristatum sub-zone fauna in the upper part of this section 
above the gasteropodous limestone is an anomaly that cannot at 


present be explained. 


4. Tor LAMELLIBRANCH FAUNA. 


The faunas collected at the localities described above consist mainly 
of Ammonites and Lamellibranchs, with comparatively few Gastro- 


298 Annals of the South African Museum. 


pods, and rare Hchinoids, Belemnites, etc. The Lamellibranchs are 
described below, and a brief notice of the associated Gastropods is 
appended. 

The assemblage of Lamellibranchs in the collections is remarkable 
for the high proportion of large, massive, and thick-shelled species, 
among which several species of Trigonia are conspicuous, e.g. T. obesa, 
T. hennigit, T. frommurzei, as well as Cucullaea spp. aff. kraussi, 
Gervillia dentata, Veniella etheridger, and Protocardia cf. sphaeroidea. 
From the systematic point of view, the chief interest in the faunas 
is the great diversity displayed by the Trigoniae, at least ten species 
of which are represented, referable to as many as seven sub-genera, 
including the very remarkable new species 7. frommurzei, for which 
the new sub-generic name Sphenotrigonia is proposed. 

A table showing the distribution of the species at the several 
localities is appended to this paper. 

Though in the last resort the precise identification of the ages of the 
several horizons must be based on the associated Ammonite fauna, 
the Lamellibranchs are of some considerable interest. On the 
Lamellibranch evidence alone, the beds would appear to range from 
Neocomian to not later than Cenomanian, and are thus largely 
Lower Cretaceousinage. The presence of the Neocomian is suggested 
by the occurrence of several species of Lamellibranchs with distinctly 
Neocomian affinities, but it appears from Dr. 8. H. Haughton’s 
preliminary analysis of the Ammonite fauna that these species are 
actually associated with Upper Aptian and even Lower Albian 
Ammonites at certain localities. The association of Trigoniae, belong- 
ing to groups hitherto regarded as exclusively Neocomian and highly 
characteristic of Neocomian deposits in Central and South America, 
the Uitenhage Formation and deposits in East Africa and India, 
with post-Neocomian ammonities, is a remarkable feature of the 
faunas. Asa result of his studies of the southern Neocomian Lamelli- 
branch faunas, Kitchin (1926, p. 467) has emphasised the fact that 
such Lamellibranch types as the Pseudo-quadrate Trigoniae, Trigoniae 
of the groups of T. conocarduformis and T. v-scripta, and large 
Cucullaeae of the C. kraussi group are confined to Neocomian deposits.* 
It is therefore somewhat surprising to find species that can be paired 
off with well-known Uitenhage (Valanginian) species from the Cape 
Province occurring in Zululand in association with Ammonites of 
later stages. Dr. Haughton finds it probable that on the Mfongosi 
“the unfossiliferous beds in the upper reaches of the spruit are Lower 

* But see p. 300. 


Lower Cretaceous Lamellubranchia from Northern Zululand. 299 


Aptian, and may range down into the Neocomian,”’ but Neocomian 
Ammonites are wanting in the collections. It is just possible that 
the Neocomian is represented on the Myesa stream (M1 and M3) and 
at ZA on the Mfongosi, since the Trigonia v-scripta group is there 
represented by a new species that was not collected at higher horizons, 
and Ammonites were not found; but in view of the undoubted 
association of e.g. a Pseudo-quadrate Trigonia with Aptian-Albian 
ammonites on the Lombagwenya and of T. obesa with Lower Albian 
ammonites at Z12, the occurrence of a new species of Trigonia belong- 
ing to the 7. v-scripta group cannot be held to prove the presence 
of the Neocomian at the localities mentioned, and it is thus possible 
that the Neocomian is not present in Zululand at all. 

Cretaceous beds older than the Middle Albian were not known 
to occur in Zululand until after the publication by Spath (1921) 
of his paper on the Cretaceous Cephalopoda of Zululand, though it 
might have been suspected that the Aptian stage, which was known 
to outcrop just north of the Zululand border, extended farther south. 
In 1929 van Hoepen published a description of the Trigoniae obtained 
by him on the Umsinene River, and in the same year claimed (1929 a, 
p. 10) that his Ndabana Beds at the base of the succession at that 
locality represented the Aptian. The evidence on which that claim 
was based has not yet been published in full. 

From his Ndabana Beds van Hoepen (1929) has only described 
four species of Trigonia, and these have a distinctly Uitenhage 
aspect, being closely comparable with species occurring in and highly 
characteristic of Neocomian deposits in Central and South America, 
and of the Uitenhage Series in the Cape Province. The occurrence 
of no less than four species of Trigonia in the Ndabana Beds that 
can all be paired off with well-known species from the Sundays 
River Beds in the Uitenhage Series of the Cape is suggestive of a 
Neocomian age for the Ndabana Beds, since at least three of these 
species belong to groups of Trigoniae highly characteristic of the 
southern Neocomian, as Kitchin (1903, 1908, 1926, 1929) has urged, 
and since also the age of the Sundays River Beds has been 
determined very precisely by Spath (1930, p. 132) as undoubtedly 
Upper Valanginian, 7.e. well below the top of the Neocomian. The 
species concerned are as follows: Megatrigonia obesa van H. belongs 
to the group of T. conocarduformis (Krauss), named after a well- 
known Uitenhage species; JLotrigonia crassitesta van H. and J. in- 
constans van H. belong to the group of 7. v-servpta Kitchin, and 
are closely comparable with T. stow: Kitchin and 7. vau Sharpe 


300 Annals of the South African Museum. 


respectively, both characteristic Uitenhage species; Pvrsotrigonia 
salebrosa van H. is a massive species of the group of the Scabrae 
distinctly reminiscent of the Uitenhage species T. kraussi Kitchin. 

The occurrence of the Neocomian in Zululand is further suggested 
by the new collections of Lamellibranchs from the Pongola area. 
In addition to T. obesa (van H.), a new species belonging to the 
T. v-scripta group (TL. haughtoni), and a large member of the group 
of the Scabrae strikingly like T. kraussa Kitchin, other character- 
istic southern Neocomian types are recorded. T. hennigi Lange is 
a very typical pseudo-quadrate Trigonia, first described from the 
T. schwarzi-Beds of the Tendaguru Formation in Tanganyika, the 
Neocomian age of which has been generally accepted (Kitchin, 1908, 
pp. 46-49; 1926, 1929), though Dietrich (1933, pp. 75-79) would 
correlate these beds with a succession ranging from Upper Valanginian 
to Lower Aptian; the species is also closely allied to two character- 
istic Uitenhage species, T. herzogi (Goldfuss) and T. holubi Kitchin, 
as well as to species in the Neocomian of South and Central America, 
and the Oomia Beds of Cutch. TZvrigonia pongolensis nov. is apparently 
very closely allied to T. krenkeli Lange from the T. schwarzi-Beds in 
Tanganyika. The massive radially costate Cucullaeae, collected on 
the Lombagwenya stream, belong to the group of which Cucullaea 
kraussi Tate from the Uitenhage Series is the type, and which has 
hitherto been regarded as confined to the southern Neocomian 
faunas, being reported also from the Neocomian of East Africa 
and of Madagascar, and from India. 

The Zululand species referred to in the preceding paragraphs are 
closely allied to species belonging to a widespread southern bivalve 
fauna which Kitchin (1926, p. 467) regarded as probably entirely 
Valanginian. The association of members of this fauna with Aptian 
and even Albian Ammonites is, however, recorded above by Dr. 
Haughton for Northern Zululand, while in India the work of Spath 
(Geol. Mag., 1935) and Cox (Pal. Indica, 1935) has shown that this 
fauna survives the Neocomian there also. 

1. Lombagwenya Spruit—The whole of the succession on the 
Lombagwenya stream (L1—L14) carries Lamellibranchs of a distinctly 
Uitenhage aspect. This is based on the occurrence of the pseudo- 
quadrate Trigonia hennigi in LY, L12, and L14, and of Cucullaea 
spp. aff. kraussi in L9 and L12. The occurrence of Panope gurgitis 
in L1, L3, L7, L9, and L11 is not inconsistent with this comparison, 
since the species ranges from Neocomian to Albian in Europe. In 
a preliminary identification of species certain Albian or post-Albian 


Lower Cretaceous Lamellibranchia from Northern Zululand. 301 


Lamellibranchs appeared to occur at these localities, but in each case 
the identification was based on doubtful material and no weight can 
therefore be attached to the record based thereon; Cucullaea woodsi 
from L12 was based on a single incomplete specimen very doubtfully 
referred to this species; <Avucellina gryphaeoides (Upper Albian to 
Turonian) from L6 was based on three rather featureless shells which 
might belong elsewhere; Hxogyra conica (Upper Albian and Cenoma- 
nian) from L8 was based on two very small specimens that might well 
belong to another species of the same genus; Pholadomya vignesi 
(Cenomanian) from L7 was based only on a small fragment; Veniella 
etheridger from L3 was based on a poorly preserved juvenile shell 
that might belong to another species of Veniella or even an unrelated 
genus; Neithea quadricostata (Albian and Cenomanian) from L5 
(Msinyene Pan) was based on a juvenile specimen referred only 
doubtiully to this species. Lopha diluviana is recorded from L3f, 
the record being based. on two specimens which belong either to 
this species or at any rate a very similar form; it is significant that 
in Kurope the species does not appear to occur before the Aptian. 

2. Myesa Spruit.—The succession on the Myesa stream (M1—M3) 
could also be compared with the Uitenhage on the Lamellibranch 
evidence, on account of the occurrence in M1 of Trigonia haughtoni and 
T. hennigi, both belonging to groups characteristic of the southern 
Neocomian, and of the former species in M3. T. obesa, which occurs 
in M1, belongs to a group known previously only from the southern 
Neocomian, but the species occurs on the Mfongosi stream as high 
up as Z12, and is apparently associated with Albian Lamellibranchs 
in Z8, Z11, and Z12. 

3. Mfongost Spruit and Pongola River.—The sequence on the 
Mfongosi stream and the Pongola. River below its junction with the 
Mfongosi appears on the Lamellibranch evidence alone to range 
from the Neocomian to the Albian, if not to the Cenomanian; the 
Senonian is absent, but whether the whole succession from the 
Neocomian to the Albian or Cenomanian is represented does not 
appear from the Lamellibranch evidence. The localities ZA, Z1-Z3 
carry Lamellibranchs suggestive of the Neocomian, the estimate of 
age being based on the occurrence of Trigonia haughtoni and T. hennigi 
in ZA, T. pongolensis in Z2, and of Cardiwm rogersi in Z2 and Z3, 
the last-named species occurring also in association with 7. haughtoni 
and T. hennigi in M1 on the Myesa. 

The horizons represented by Z4—Z14 are not certain from the 
Lamellibranch evidence. The occurrence of Trigonia obesa in Z4, 

VOL. XXXI, PART 3. 26 


302 Annals of the South African Museum. 


Z5, Z8, and Z12, and of a large Scabroid Trigonia comparable with 
T. kraussi in Z8, is suggestive of a Neocomian age for these horizons, 
since these species belong to groups of Trigonia characteristic of 
southern Neocomian deposits.* On the other hand, a preliminary 
identification of species gave certain Albian species as occurring in 
Z6, Z8, Z9, Z11-Z14. Among the latter is Cucullaea woods, described 
by Newton from the Manuan Creek area, and said by du Toit (1926, 
p- 318) to come from the upper portion of the Manuan Creek (Albian). 
The record of Glycymeris griesbachi (Albian and Cenomanian at 
Manuan Creek, according to du Toit) from Z8 was based on a single and 
not very well preserved specimen, and may be incorrect. The record 
of Exogyra conica (Upper Albian and Cenomanian) from Z8 was based 
on one small shell, and it is possible that it belongs to another species. 
The record of Neithea quadricostata (Albian and Cenomanian) from Z9 
was based on an extremely unsatisfactory specimen, an incomplete 
and juvenile shell. Nethea quinquecostata, recorded from Z13, is 
very long ranged, having been recorded from the Aptian to the 
Senonian. Inoceramus concentricus, a characteristic Upper Albian 
species in Hurope, was apparently satisfactorily determined from Z6 
and Z14, but in each case the record is based on one specimen. In 
determining the horizons of Z4—Z12 the only Lamellibranch evidence 
which appears to be of possible significance is the occurrence of 
T. obesa at several localities (Z4, Z5, Z8, and Z12), the presumed 
occurrence of the Albian species I. concentricus in Z6, and the fact 
that C. woodsi, which occurs in Z8, Z11, and Z12, occurs also ina 
typical Albian association in Z19. The horizons present at these 
localities must be determined on the evidence of the Ammonites. 

The localities north of the Mfongosi and below its junction with 
the Pongola River (Z16—-Z21) are, on the Lamellibranch evidence, 
either Albian or Albian and Cenomanian, no higher horizon being 
represented. The correlation is based on the occurrence of a number 
of species which occur in the Manuan Creek area, some of which 
occur in the Albian and Cenomanian of Europe and elsewhere. 
Pholadomya vignesi occurs in the Cenomanian in Syria, but is said 
by du Toit (1926, p. 318) to occur in the Albian of the Manuan 
Creek, while it makes its appearance in Z16 (7 typical specimens), 
and appears also in Z19-Z21. Ezxogyra conica (Upper Albian and 
Cenomanian) occurs in Z19 and Z20. Neithea quadricostata (Albian 
and Cenomanian) is represented by fine specimens in Z19 and Z20. 
Inoceramus concentricus (Upper Albian) occurs in Z19, and its variety 

* But see p. 300. 


Lower Cretaceous Lamellibranchia from Northern Zululand. 303 


subsulcatus in Z16, Z19, Z21. In addition to the above, several 
species occur which have been described from the Albian-Cenomanian 
succession on the Manuan Creek and the Umsinene River, Veniella 
etheridgea and the Gastropod Turritella manuanensis making their 
appearance in Z16. 

4. Mokatuni.—The species recorded from Z25, south-east of Moka- 
tini and south of the Pongola River, include Trigonia blanckenhorni, 
Trigonarca ci. lgeriensis, Veniella etheridgei, and probably also 
Exogyra conica (one flat valve), which is very similar to the assemblage 
recorded by Newton (1909, p. 88) from the north end of False Bay 
(Zululand), probably Cenomanian in age. 

Note on certain Lamellibranchs from the Uitenhage Series.—This 
opportunity is taken of publishing additional figures of certain 
Uitenhage species of Lamellibranchs described originally by Krauss 
(1850), Tate (1867), and Neumayr (1881), but which were not re- 
figured by Kitchin (1908) in his monograph on the fauna of the 
Uitenhage Series, as well as figures of an adult individual of Trigonia 
conocarduformis. Additional information with regard to these species 
is forthcoming, or additional figures are required. These species are 
referred to in the systematic descriptions of Zululand shells which 
follow, and only passing reference need be made here. The species 
concerned are as follows:— 

Cucullaea kraussi Tate (see p. 305, Plate XLIV, fig. 2). Additional 
figures are necessary to show the finer radial ribbing present in 
addition to the coarse radial costae. The species becomes the type 
of the new sub-genus Megacucullaea. 

Gervillia dentata Krauss (see p. 310, Plates LIII-LV), the original 
figures of Krauss are insufficient to show the form of the species. 

Trigonia conocardiiformis (Krauss) (see p. 337, Plate XL, figs. 
1-3); the species was well described by Kitchin, but only juvenile 
and half-grown specimens were figured. A fine adult shell (Alb. 
Mus. 788) is here figured for comparison with 7. obesa. 

“Anoplomya” lutraria Krauss (see p. 385, Plate L, figs. 2-4, 
7, 10, 11); this species is probably in the synonymy of Panope 
gurgitis. Additional figures are given for comparison with shells from 
Northern Zululand. 


304 Annals of the South African Museum. 


5. DESCRIPTION OF THE SPECIES. 
Famity ARCIDAE. 
Genus CucuLLAEA Lamarck 1801. 


Sub-genus Cyphowis Rafinesque 1819. _ 
Cucullaea (Cyphoxis) woodsi R. B. Newton. 


1909. C. woodst Newton: Trans. Roy. Soc. 8. Afr., vol. i, p. 31, 
pl. iv, figs. 4-9. 

This species occupies a unique position in the South African faunas, 
and has been sufficiently well described by Newton. Its occurrence 
on the Mfongosi stream and neighbourhood is as follows: Z8 (2, 
typical), Z11 (8, doubtful), Z12 (1), Z19 (35, fine and very typical 
set). The species was described with a mixed Cretaceous fauna from 
the Manuan Creek in Zululand, but is said to be from the Albian 
at that locality (du Toit, 1926, p. 318). As is pointed out below, 
the typical Cretaceous Cucullaeae should be referred to Cyphozis 
Rafinesque. 


Cucullaea Lamarck 1801, [donearca Conrad 1862, and Cyphozis 
Rafinesque 1819. 


According to Gardner (1916, p. 529) and Stewart (1930, p. 74) the 
genus Cucullaea was founded by Lamarck in 1801 (Syst. Anim. sans 
Vert., p. 116), and the genotype, subsequently designated by Children 
(Lam. Gen. Shells, 1823, p. 45), is the recent Oriental species Cucullaea 
auriculifera Lamarck, for which the earliest name appears to be 
Arca concamerata Martini, 1777. Stewart states that the Mesozoic 
species commonly included in Cucullaea not only have a much 
thicker shell than the genotype, but also have diagonal grooves on 
the ligamental area which are lacking in the latter. According to 
him it does not seem that the Mesozoic group can be regarded as 
more than a sub-genus. 

The generic name [donearca proposed by Conrad in 1862 (Pr. Ac. 
Nat. Sci. Philad., p. 289) for the Cretaceous species C. tippana Conrad, 
has been widely used for the Mesozoic Cucullaeae. The type species 
is usually placed in the synonymy of C. vulgaris Morton (Gardner, 
1916, p. 529, pl. xx, figs. 8,9; pl. xxi, figs. 1, 2; Wade, 1926, p. 43, 


Lower Cretaceous Lamellibranchia from Northern Zululand. 305 


_ pil. ix, figs. 3, 4, 6, 7), one of the most common Lamellibranchs in the 
Upper Cretaceous of the eastern United States. On the other hand, 
Gardner, Wade, and Dietrich (1933, p. 28) have sunk Idonearca in 
Cucullaea, though Gillet (1924, Lamell. Néocom., Mém. Soc. Géol. 
France) has maintained the distinction. The differences noted by 
Stewart, together with the fact that the Mesozoic type apparently 
failed to survive the Lower EHocene, seem sufficient to warrant the 
retention of a distinct sub-genus for the Mesozoic species. 

Stewart (1930, p. 75) states that Pilsbry has found that the name 
Cyphoxis Rafinesque 1819 (Jr. de Physique, vol. 88, p. 427) is an 
earlier name for Cretaceous Cucullaeae, and has accordingly referred 
Gabb’s species to the sub-genus Cyphowis. 


Sub-genus Megacucullaea sub-gen. nov. 


Type.—C. kraussi Tate. 

The large, massive, radially costate South African Neocomian 
species Cucullaea kraussi Tate appears to be sufficiently distinct from 
the recent Cucullaea and from Cyphoxis (=Idonearca) for it to be 
placed in a separate sub-genus. The species was originally described 
by Krauss (1850, p. 452, taf. 48, fig. 2) as C. cancelllata, the name 
being later changed by Tate (1867, p. 161) to C. kraussi as the former 
name was preoccupied. The species was well described by Neumayr 
(1881, p. 275, taf. 2, fig. 2), who referred to its isolation as a species 
and to the difference between it and Cucullaea (sensu stricto), but the 
species, which is by no means rare in collections from the Sundays 
River Beds of the Uitenhage Series, does not appear to have been 
figured since in a palaeontological publication. According to Dietrich 
(1933, p. 28) the species was placed by Gillet (Lamell. Néocom., 
1924) in the genus Idonearca, but in a “section spéciale’”’ on account 
of its peculiar massiveness and strong ribs. 

Similar forms have been recorded from Hast Africa (Lange, 1914, 
p- 223; Dietrich, 1933, p. 28) as C. kraussi, but these do not appear 
to have been figured. Kitchin (1908, p. 43) has mentioned the 
occurrence of fragments of a similar Cucullaea in the Oomia Beds 
in India, and Dietrich (loc. cit.) states that Gillet has recorded the 
species from Madagascar. Dietrich also states that C. neuquensis 
Weaver (1931, p. 186) from the Argentine belongs to the same group. 
The sub-genus appears to be a characteristic element of the southern 
Neocomian fauna, and is here recorded also from post-Neocomian 


beds in Zululand. 


306 Annals of the South African Museum. 


The genotype 1s characterised by the following salient features: 
Shell very large, presumably equivalve, more or less equilateral, 
roughly triangular in outline, very strongly inflated, longer than 
high. Shell substance very thick. Anterior and ventral margins 
rounded, forming a continuous broad curve, coarsely scalloped. 
Posterior margin relatively short, more or less straight, sloping 
steeply downwards, and meeting the ventral margin in a rounded 
postero-ventral angle. A strong, rounded posterior carina, sepa- 
rating a steep concave posterior area from the rest of the valve, 
persists from the umbo to the postero-ventral angle. Umbonal 
region broad, very prominent, more or less centrally placed; umbo 
very strongly incurved, free and projecting, at some distance from 
the hinge margin, showing very little forward curvature. Hinge 
line long, straight, a little shorter than the greatest length of the 
shell. Hinge very narrow except at the two ends, with numerous 
small transverse teeth for more than half of its length, the central 
teeth short and vertical, passing laterally into slightly longer oblique 
teeth; at each end there are three considerably larger teeth approxi- 
mately parallel to the hinge margin, the upper two long, the lower 
shorter. Area very large, remarkably wide, triangular, concave; 
area with numerous (15-18) deeply incised ligament grooves, sepa- 
rated by smooth, flat interspaces and traversing the area more or 
less parallel to its upper margin. Interior of valves smooth, faintly 
undulate near the ventral border; muscle impressions large, the 
posterior a bit sunken but without a raised plate. 

Shell surface ornamented with very strong radial costae, numerous 
radial riblets, and with growth lines. Between the posterior carina 
and the anterior slope there are several (7 or 8) very strong, rela- 
tively sharp costae, triangular in cross-section, separated by broad 
concave interspaces. In addition there may be a weaker rib in the 
broad space between the carina and the first of the costae, as well 
as 2 or 3 weaker ribs on the anterior slope; the posterior area has 
2 or 3 weak ribs. Radial riblets very numerous, rounded, separated 
by rounded interspaces, varying very much in strength, present on 
all parts of the valve, but tending to be absent from the vicinity 
of the hinge margin; with full maturity the riblets die out before 
reaching the ventral border. Growth lines and growth lamellae 
throughout, tending to form a lattice with the riblets, and giving 
rise to notches on the crests of the costae. 

The most striking characters of the sub-genus, which serve to dis- 
tinguish it even at a glance from the living genotype of Cucullaea 


Lower Cretaceous Lamellibranchia from Northern Zululand. 307 


and from Cyphoxis (=Idonearca), are the very large size, the extra- 
ordinary degree of inflation, the very wide area with very numerous 
grooves, and the prominent radial costae. The genotype of Cucullaea 
is said by Stewart (1930, p. 75) to be thin and without grooves on 
the area, and judging from Fischer (1887, p. 977, pl. xvii, fig. 14) 
is a smaller shell with numerous radial riblets, a ventral border very 
finely scalloped within, a relatively narrow area, a raised plate for 
the attachment of the posterior adductor muscle, and a less differ- 
entiated dentition with the teeth radially arranged. C. kraussi 
certainly cannot be included in Cucullaea (sensu stricto) in view of 
the differences above noted, the absence of the raised plate for 
the posterior adductor (“an essential character of Cucullaea’’— 
Arkell, 1930, p. 309), the strongly developed costation and the 
ponderous form being of importance. The dentition is also different, 
for in C. kraussi the short vertical teeth at the centre of the hinge 
are transitional into longer oblique teeth which are directed laterally 
downwards, and there is an abrupt change when the horizontal teeth 
begin; in the genotype of Cucullaea the teeth are directed radially 
upwards, and the shorter central teeth pass gradually into the rather 
longer very oblique lateral teeth. 

I am not acquainted with the genotype of Cyphowis, but the species 
presumably has the. same general features as the genotype of 
Idonearca, C. vulgaris Morton (vide supra). The latter is a very 
characteristic Cretaceous Cucullaea and is distinguished from C. 
kraussi in several important respects. In the first place C. kraussi 
is distinguished by its larger size, the great thickness of the shell 
substance, the extraordinary degree of inflation, the great prominence 
of the umbonal region, the great width of the area, and the very 
large number of ligament grooves. In the second place C. kraussi 
is distinguished by the very prominent radial costae which have no 
counterpart in C. vulgaris; the radial costae are present in addition 
to the much finer, more or less reticulate, ornament of radial riblets 
and concentric growth lines characteristic of C. vulgaris and other 
species; the costae are correlated with the scalloping of the ventral 
margin, which in C. vulgaris is smooth. In the third place the 
posterior adductor impression of C. kraussi is, according to Neumayr, 
without the raised plate or radial buttress which occurs in C. vulgaris 
and the Cucullaeae generally. In the fourth place C. kraussi is 
distinguished by the relatively smaller teeth, the central teeth being 
relatively shorter and more than twice as numerous as in C. vulgaris. 
In view of these striking differences the exclusion of C. kraussi from 


308 Annals of the South African Museum. 


Idonearca, where it was placed by Gillet, is imperative, and a new 
sub-generic name is proposed. 

C. kraussi has not been adequately described or figured, and the 
opportunity is here taken of publishing an additionalfigure (Plate XLIV, 
fig. 2) and of emending the descriptions given by Krauss and Neumayr 
in the publications cited above. The species was founded by Krauss 
on a single incomplete left valve, and Neumayr’s fine figures were 
based on a complete left valve, the only specimen in the Holub 
collection. The original specimen apparently showed no traces of 
radial riblets, for these were neither mentioned by Krauss nor de- 
picted in the figure; the broad interspaces between the costae were 
said to be crossed by growth lines which, in groups of 4-8, gave rise 
to alternate raised bands and furrows, 7.e. a concentric undulation 
of the valve surface, giving the surface a latticed appearance. 
Neumayr noted only in one place on his specimen a fine radial 
striation, which formed a lattice with the growth lines, which he 
merely described as strong and somewhat scaly. Well-preserved 
specimens show that radial riblets, numerous and rounded, are a 
characteristic feature of the greater part of the shell surface, and 
these have been described above and are here depicted on the ac- 
companying figure; their absence or rarity in the earlier described 
specimens is presumably due to their condition of preservation. 
The concentric undulation of the surface is not usually as regular 
as depicted by Krauss, and becomes very indistinct towards the 
ventral border of nature shells. 


Cucullaea (Megacucullaea) spp. indet. 
(Plate XX XVII, figs. 1-3.) 


The sub-genus is represented in the collection made by Dr. 8. H. 
Haughton and Dr. A. W. Rogers by three very imperfect shells 
having a close general resemblance to C. kraussi Tate and undoubtedly 
closely allied to that species. The specimens, unfortunately, are too 
incomplete, weathered, or crushed for a satisfactory comparison. The 
occurrence of the sub-genus in Zululand is, however, of some interest. 

The specimens were collected at two localities on the Lombagwenya 
tributary of the Pongola River, in both cases in association with 
Trigonia (Steenmanella) hennigr Lange: at L9 two fragments of the 
anterior part of a left valve; at L12 a complete but very weathered 
and crushed left valve, as well as another weathered left valve 
preserved partly as an internal cast. 


Lower Cretaceous Lamellibranchia from Northern Zululand. 309 


The fragments from L9 are portions of a very large mature, highly 
inflated, very thick shell, showing a small portion of the area, the 
terminations of the anterior costae, and a broad marginal band with 
growth lines only. Finer radial ornament is not preserved, but the 
fragments could well be portions of a shell of C. kraussi. The area 
is traversed by very numerous parallel ridges and grooves resembling 
ligament grooves, but these have been produced by weathering, and 
traces of the more widely spaced ligament grooves are preserved, 
crossing the former obliquely. | 

The more complete shell from L12 is at first sight rather different 
from C. krausst by reason of the fact that the umbonal region is 
more prominent and the apical angle much less than in that species. 
An examination shows, however, that the specimen has been badly 
crushed, so that the anterior slope has been considerably increased 
and the incurved tip of the umbo pressed down on to the area. To 
what extent the difference in form noted is original or due to crush- 
ing I could not satisfactorily determine. The specimen is badly 
weathered and not only shows no sign of fine radial ornament 
but only traces of the growth lines. In front of the posterior 
carina there are six prominent costae, as in C. kraussi, and these 
appear to have been of the same character as those of the 
Uitenhage species. -On the posterior slope there are four finer 
radial ribs. 

The second left valve from L12, though incomplete, seems to be 
distinct from C. kraussi. The specimen is weathered and the thick 
shell is preserved only in the anterior third and on the posterior slope, 
the central portion being in the condition of an internal cast. The 
cast shows faint, broad, radial undulations and the impression of a 
narrow, raised, internal rib, which proceeds almost vertically from the 
posterior side of the umbo for a distance of 30 mm. The general 
form is identical with that of a medium-sized C. kraussi and the 
posterior slope is similarly ornamented. The radial costae seen at 
the antero-ventral margin differ from those of C. kraussi, in being 
conspicuously broader; the two uppermost costae at the anterior 
end are narrow and raised as in C. kraussi, but the next three costae 
are broad and convex and carry on their summits a distinctly de- 
marcated narrow rib or crest; in one case there is a trace of a sub- 
sidiary rib on the flank; these costae are followed by a smaller 
strong rib. The interspaces are more or less deeply U-shaped, are 
crossed by growth lines, and show traces of radial riblets. The 
difference in costation is so marked that a distinct species might well 


310 Annals of the South African Museum. 


be represented, but the specimen is too poor to be made the holotype 
of a new species. 


Genus TrigonaRrca Conrad 1862 
(= TrIGonoARcA Conrad 1867). 
Trigonarca cf. ligeriensis (d’Orbigny). 


1909. T. lagerrensis (d’Orb.): R. B. Newton, Trans. Roy. Soc. S. 
Afr., vol. 1, p. 33, pl. iv, figs. 13-18. 


Newton figured three specimens, referred to this typically Ceno- 
manian species, from the north end of False Bay in Zululand, where 
it occurs with Trigonia blanckenhorni and Exogyra conica. A single 
imperfect specimen that resembles the above was found in the same 
association south of Mokatini, Z25. 

The original spelling of the generic name is Trigonarca (fide 
Stephenson, 1923, p. 96), and for form Trigonoarca used by Woods 
(1906, p. 288) and Rennie (1930, pp. 169, 240) for South African 
species 1s incorrect. 


Genus GLYCYMERIS da Costa 1778. 
Glycymeris cf. griesbacht R. B. Newton. 
1909. G. griesbacht Newton: Trans. Roy. Soc. S. Afr., vol. i, p. 36, 
pl. i, figs. 13-17 (non G. griesbachi Etheridge, Saxicavidae). 


The type set 1s from the north end of False Bay in Zululand, where 
the species occurs with Hxogyra conica (J. Sow.). The occurrence is 
as follows, though in every case the shells are imperfectly preserved 
and the identification is somewhat uncertain: Mfongosi stream 
Z8 (1); Pongola River below junction with Mfongosi, Z19 (4). 


Famity ISOGNOMONIDAE (=PERNIDAB). 


Genus GERVILLIA Defrance 1820. 
Gervillia dentata Krauss. 


(Pl. Li ngs: 1525 Pl. Lilies: 1-35) 
(Pls. LILI-LV, from the Uitenhage Series.) 


1850. G. dentata Krauss: Untere Kreide Kaplandes, p. 458, pl. 1, 
figs. 1 a-c. 


Lower Cretaceous Lamellibranchia from Northern Zululand. 311 


11907. G. dentata Krauss: Etheridge, Rep. Geol. Natal, 3, p. 78, 
pl. 1, figs. 13-15. 

1908. G. dentata Krauss: Kitchin, Ann. 8. Afr. Mus., vol. vii, p. 36. 

1909. G. sublanceolata (d’Orb.): Newton, Trans. Roy. Soc. 8S. Afr., 
vol. i, p. 48, pl. in, figs. 7-10. 


Material.—Large Gervilliae similar to G. dentata Krauss from the 
Uitenhage formation are of frequent occurrence in the Cretaceous 
of Northern Zululand. Dr. 8. H. Haughton and Dr. A. W. Rogers 
collected specimens from the following horizons along the Mfongosi 
stream and neighbourhood, the number of examples being indicated 
in brackets: Z3 (1), Z4 (3), Z6 (3), Z7 (7), Z8 (10), Z10 (2), Z16 (1), 
Z19 (4). Mr. H. F. Frommurze collected a number of specimens 
at the same locality: S.A.M. 10802-10807; whilst Mr. J. 8S. Hutt 
collected the specimens numbered T.M. 1661-1666, 1668, 1670-1, 
1673, 1678-9, 1682, 1684, 1685, 1687, 1911.* 

Remarks.—Krauss figured three examples of this species from the 
Sundays River Beds of the Uitenhage Series, but the species has not 
been described by subsequent authors, although examples are not 
infrequent in collections. The species has a close general resemb- 
lance to the Huropean G. sublanceolata (d’Orb.) and G. anceps 
Deshayes. R. B. Newton (1909, p. 48, pl. iu, figs. 7-10) described 
some rather imperfect shells from the north end of False Bay in 
Zululand, associated with ammonites identified as Cenomanian by 
Crick (1907), as G. sublanceolata, and stated that G. dentata is dis- 
tinguished by its more convex and more robust valves and by the 
fact that the antero-ventral margin is not concave. With a repre- 
sentative set of G. dentata before me (see Plates LIII-LV), including 
some examples from the cliffs opposite Redhouse on the Zwartkops 
River (the vicinity of the locality from which came the type specimens), 
I have found great difficulty in drawing any distinction between that 
species and the so-called G. sublanceolata from False Bay, and it may 
well be that these, together with the new specimens from the Mfongosi, 
belong to G. dentata. 

The figures given by Krauss show the anterior part of the ventral 
margin very slightly convex, and this point has been used by Newton 
in drawing a distinction between G. dentata and G. sublanceolata, 
though the distinction is of no great magnitude. In fact, the anterior 
part of the ventral margin is very nearly straight in all the forms here 


* The species was also recorded by Dr. S. H. Haughton at the following 
localities: Z5, L3, L12, L4, L6, Ll. 


312 Annals of the South African Museum. 


being considered. In the specimens from the Uitenhage Series before 
me the anterior part of the ventral margin is either very nearly straight 
or quite distinctly but gently concave, the latter condition being by 
no means uncommon. In this respect some of the Uitenhage shells 
agree exactly with the forms figured by Newton from False Bay, as 
well as with Woods’s figures of G. sublanceolata from the Lower and 
Upper Greensands (1905, p. 74, pl. x, figs. 14-16; pl. x1, fig. 1; text- 
figs. 7 a—c, 8). A very gentle concavity is usually discernible among 
the Mfongosi specimens also. Since it can scarcely be doubted that 
the specimens before me from Redhouse are examples of Krauss’s 
species, it is clear that the distinction based on this feature must 
be dropped. 

Newton further states that G. dentata is of greater thickness, more 
robust, and more convex than G. sublanceolata. There is little in 
the figures given by Krauss to warrant this statement, though it 
would appear that the shell substance is by no means thin in the 
former. The shells before me show quite a considerable range of 
variation in respect to massiveness, including both somewhat convex 
and more flattened forms; the shell substance in one instance is as 
much as 13 mm. thick in the middle part, though usually very con- 
siderably less, so that the valves can in some instances be described 
as relatively thin. 

The shells figured by Newton from False Bay were found in associa- 
tion with Hxogyra conica (J. Sow.), and with Glycimeris griesbachi 
Newton (Newton, 1909, p. 88), and it is significant that the same 
association is encountered in Z8 and Z19 on the Mfongosi, though 
Gervilliae extend down into beds of possible Neocomian age at that 
locality. The specimens from the Mfongosi are in general agreement 
with those figured by Newton, and must be the same species. The 
former have been carefully compared with examples of G. dentata, 
and the comparison has failed to reveal any general distinction, 
though in one or two instances it was noticed that the apical angle 
was somewhat larger than is the case with either the Uitenhage or 
Lower Greensand shells, and consequently the posterior wing was 
relatively broader; this latter condition is, however, by no means 
general, and in the majority of specimens no distinction can be drawn 
in this respect. 

A number of specimens from Uitenhage and the Mfongosi would 
appear to be scarcely distinguishable from the specimens of G. sub- 
lanceolata figured by Woods, the agreement extending to the details 
of form used in distinguishing the species from its European allies, 


Lower Cretaceous Lamellibranchia from Northern Zululand. 313 


the slightly concave antero-ventral margin, the central position of 
the line of greatest convexity, and the nearly straight posterior 
margin of the posterior wing, and extending also to the nature and 
number of the ligament pits and to the dentition. It should be 
pointed out, however, that G. sublanceolata is apparently very closely 
allied to certain other Huropean species, in particular to G. anceps 
Deshayes, with which it has frequently been confused, and that a 
satisfactory comparison with G. dentata cannot be made without an 
actual examination of representative sets of these European species. 
Only in one particular does there seem to be a constant point 
of difference between the Uitenhage shells before me and those from 
Atherfield figured by Woods. In the latter the left valve is distin- 
guished from the right by having a demarcated, antero-ventral area 
below the pointed umbo; this area is narrow, elongate, and bounded 
by a linear depression, its length less than one-third of the total 
length of the valve. In the Uitenhage shells the same distinction 
between the valves is evident, but the antero-ventral area in the left 
valve may be of considerable length and width, extending backwards 
for more than one-third of the total length of the valve, in one instance 
nearly one-half of the length of the valve. The length and width 
of this demarcated area varies considerably, though in every instance 
the contrast with the much smaller feature in the Atherfield shells 
is noteworthy. The area is convex and is distinguished only in that 
it is bounded by an impressed line, and in that there is a discrepancy 
between the growth lines of area and flank. Newton does not mention 
the feature in the case of the False Bay shells, and the figured left 
valve was probably too poorly preserved to show it. The shells from 
the Mfongosi are in general somewhat weathered, and the feature 
is not easily discerned on that account, though in one or two instances 
it is apparently indicated, the area in question being relatively wide 
and persisting for about one-third of the total length of the shell, 
the shell exactly matching a similar valve from the Uitenhage Series. 
The distinctions drawn by Newton between G. dentata and G. sub- 
lanceolata have been shown to be without substance, and if a distinc- 
tion is to be maintained between these species the difference in the 
size of the demarcated antero-ventral area will have to be relied on. 
In every other respect the species would appear to be very similar, and 
it is significant that Newton, who presumably was able to compare 
directly the shells from False Bay and Atherfield, did not hesitate 
to identify the False Bay shells with G. sublanceolata. Certain 
specimens from the Uitenhage Series have this antero-ventral area 


314 Annals of the South African Museum. 


so distinctly larger than in the shells from Atherfield, that a specific 
distinction might well be maintained in spite of general agreement 
in other respects. The Mfongosi shells appear to agree with certain 
examples of G. dentata in respect to this feature, and to them and the 
shells from False Bay the name G. dentata might well be applied. 
It must be admitted, however, that the state of preservation of the 
Zululand material leaves much to be desired and the comparison 
is not ultimately satisfactory. 

The specimens figured by Etheridge (loc. cit.) from the Umsinene 
River in Zululand, though presumably the same species as the other 
Zululand shells, are too imperfect for fine distinctions to be drawn. 


Genus Isognomon Solander 1786 
(= Perna Bruguiére 1789). 
Isogonomon sp. 


Rather imperfect specimens of a large species, sometimes showing 
the ligament pits, were recovered from the following localities: 
Lombagwenya stream, L3f (1); Mfongosi stream, Z7 (1), Z8 (1), 
Z14 (2). 


Genus INocERAMUS Parkinson 1819. 
Inoceramus concentricus Parkinson. 


1910. I. concentricus Park.: Woods, Cret. Lam. England, vol. ii, 
p. 265, pls. xlv—xlvu. 

1930. I. concentricus Park.: Heinz, Ueber Inoceramen Siidafrika, 
Comp. Rendu, 15th Int. Geol. Congress, vol. 1, p. 683, 
fens 


Heinz collected a single specimen of this well-known species at 
the Manuan Creek (Munyuana) from an horizon identified by van 
Hoepen as Albian. According to Heinz the species is a key-fossil 
of the Upper Gault in Europe. What is almost certainly the same 
species was collected on the Mfongosi stream and neighbourhood: 
Z6 (1), Z14 (1), Z19 (1). 


Inoceramus concentricus Park. var. subsulcatus Wiltshire. 


1910. I. concentricus var. subsulcatus Wilts.: Woods, Cret. Lam. 
England, vol. 1, p. 268; pl. xlvu, figs. 3-14. 


Lower Cretaceous Lamellibranchia from Northern Zululand. 315 


A radially sulcate Inoceramus, which appears to be identical with 
the form described by Woods, is sparingly represented in the collection 
from the neighbourhood of the Mfongosi stream: Z16 (2), Z19 (1), 
Z21 (1). The form has not hitherto been recorded from South 
Africa, though in the collection of the Transvaal Museum there is 
a well-preserved specimen (T.M. 1284) in an Albian fauna from 
Catuane, a locality some thirty miles distant on the southern border 
of Portuguese Hast Africa. 


Famity PTERIIDAE. 
Genus PsEupavicuLa R. Etheridge, fil., 1892. 
Pseudavicula? africana R. Etheridge, fil. 


1907. Pseudavicula? africana R. Eth., fil.: 3rd Rep. Geol. Natal, 
p. 71, pl. xi, figs. 8-11. 

Small shells similar to those named by Etheridge from the Umsinene 
River in Zululand, and referred by him very doubtfully to Pseudavicula, 
are very abundant at certain horizons on the Mfongosi, mostly in 
an exfoliated condition; these were collected as follows: Z4 (abundant), 
Z5 (4), Z11 (5), Z13 (several). The same species was also collected 
at Z24 (abundant) south of Mokatini. 


Famity MYALINIDAE. 


Genus AUCELLINA Pompeckj 1901. 
Aucellina gryphaeoides (J. de C. Sowerby) ?. 


1905. Aucellina gryphaeoides (J. de C. Sow.): Woods, Cret. Lam. 
England, vol. u, p. 72, pl. x, figs. 6-13. 


In England this species ranges from Upper Albian to the top 
of the Cenomanian. Shells referred doubtfully to the species were 
collected on the Lombagwenya stream at L6 (3). At Catuane on 
the Zululand-Portuguese Hast African border the species is present 
in an Albian fauna preserved in the Transvaal Museum, some of the 
specimens being very finely preserved (10 specimens, T.M. 1257, 1258, 
1265-7, 1295). 


316 Annals of the South African Museum. 


Famity OSTREIDAE. 
Genus Exocyra Say 1820. 


Exogyra conica (J. Sowerby). 


1909. Exogyra conica (J. Sow.): R. B. Newton, Trans. Roy. Soc. 
S. Afr., vol..i, p,:51, pl. xi,.figs. 8-10, 

1912. Exogyra conica (J. Sow.): Woods, Cret. Lam. England, vol. 
ii, p. 407, figs. 215-242. 

Newton figured specimens from the north end of False Bay and 
from the Manuan Creek in Zululand. The species is said to be Upper 
Albian and Cenomanian. In the present collections there are 
examples from the following: Lombagwenya stream, L8 (2, small ?); 
Mfongosi stream, Z8 (1, small?); Pongola River below junction with 
Mfongosi, Z19 (15, fine set), Z20 (2); south of Mokatini, Z25 (1, flat 
or right valve, presumably the same species). 

The species was also recorded by Dr. 8. H. Haughton at lonalate 
Z16 and L6. 

Genus LopHa Bolten 1798 


(= ALEctTRyoni4 Fischer de Waldheim 1807). 
Lopha diluviana (Linn.). 


1912. Ostrea diluriana Linn. Woods: Cret. Lam. England, vol. ii, 
p. 342, figs. 98-138. 


This species, which in Europe ranges from the Aptian to the Lower 
Chalk, is apparently represented in the collections from L3} and 
Z19. The specimens from L37 (2) on the Lombagwenya stream are 
of the characteristic elongate type and agree very closely with the 
English examples figured by Woods. Those from Z19 (4) on the 
Pongola River below its junction with the Mfongosi stream are short 
and coarse ribbed, and are referred somewhat doubtfully to the 
species; they are not unlike Ostrea milletiana d’Orbigny from the 
Gault, a species included in the synonymy of O. diluviana by Woods. 


Famity TRIGONIIDAE. 
Genus Triconia (Bruguiére 1789) Lamarck 1799. 


Genotype T. sulcata (Hermann, § Costatae). 


During the past decade a number of authors (Deecke, 1925; van 
Hoepen, 1929; Crickmay, 1930, 1930 a; Marwick, 1932; Dietrich, 


Lower Cretaceous Lamellibranchia from Northern Zululand. 317 


1933) have proposed new generic or sub-generic terms for species of 
Trigoma from Jurassic and Cretaceous formations. Several of these 
are founded on or are applicable to the species described below. 
In view of the divergent views recently expressed as to the genotype 
of Trigonia, and in view also of the fact that a considerable number 
of genera, grouped in four sub-families, were proposed for Zululand 
species by van Hoepen (1929), and the present writer’s preference 
for accepting these terms as of not more than sub-generic value, a 
general statement of his position will not be out of place. 

The Genotype of Trigonia.—The genus was founded by Bruguiére 
(1789, Encycl. Méth. Vers, vol. i, p. xiv) with short diagnosis but 
without mention of species by name, without figures or references. 
His diagnosis reads: “‘Coquille triangulaire inéquilatérale, charniére 
composée de deux grosses dents plats, trés saillantes et sillonées des 
deux cotés.” According to Article 25 a* of the International Rules 
of Nomenclature this is sufficient to found the genus if the author 
has applied the principles of binary nomenclature, which Bruguiere 
certainly has done elsewhere in the same volume. Such a case 
appears to be covered by Opinion 46,+ the last part of which reads: 
“Tf it is not evident from the original publication of the genus how 
many or what species are involved, the genus contains all the species 
of the world which would come under the generic description as 
originally published, and the first species published in connection 
with the genus becomes zpso facto the type.” From this it would 
follow that Trigoma Bruguiére 1789 is valid, and indeed the validity 
has seldom been questioned. The Rules are ambiguous and the 
various Opinions contradictory, however, since it might be held that 
the whole spirit of Article 30 (designation of type species) requires 
the genus to be accompanied by a named species; ¢ if this view is 
adopted then Trigonia would have to be cited from the earliest 
reference of a named species to the genus. 

The name appeared again in 1797 at the top of Plates 237 and 
238 § of the Tabl. Encycl. Méth. Vers, on which unnamed species of 
Trigonia were figured by Bruguiére. Plate 237 consists entirely of 
shells which we should refer to Trigonia (sensu lato), but Plate 238 


* [Xe Congrés Internat. Zool. 4 Monaco (Rennes, 1914), p. 901. 

7 Ibid., p. 914. 

+ I am indebted to Dr. K. H. Barnard for pointing this out to me. 

§ Stewart (1930, p. 88) and others have failed to mention that the name Trigonia 
heads two plates, and we are indebted to Crickmay (1932, p. 446) for pointing out 
that the name appears on Plate 238 as well as on Plate 237. 

VOL. XXXI, PART 3. 27 


318 Annals of the South African Museum. 


includes in figs. 3a,b, one species which we should refer to Lima 
(Plagiostoma) and which would not fall under the diagnosis given 
by Bruguiére in 1789. It may be doubted whether species figured 
under a generic name but not named specifically have any claim to 
be considered as genosyntypes to the exclusion of all other species. 
In the present instance two courses have to be taken into account: 
(1) the genosyntypes are the unnamed species figured on Plates 237 
and 238 of Tabl. Encycl. Méth. Vers, Bruguiére 1797, and either the 
first specific name to be applied to any of these figures becomes the 
type by monotypy, or it is competent for a subsequent author to 
designate a genotype from among the several specific names applied 
to these figures in the interim; (2) the unnamed species figured on 
Plates 237 and 238 are not necessarily the genosyntypes, and since 
it is competent for a subsequent author to refer to the genus any 
named species which would fall under the diagnosis given by 
Bruguiere 1789, the genotype would then be either the first-named 
species so referred or among the several named species so referred. 

Stewart (1930, p. 88) has adopted the former course, for he rejects 
T. pectinata Lam. on the grounds that it 1s not available, though he 
erroneously restricts the genosyntypes to the figures on the first of 
the two plates. The figures on Plate 237 were later given names 
by Lamarck, and one of these has usually been cited as genotype. 
The names given were:* TJ. nodulosa Lam. for fig. 4 in 1801 (Syst. 
Anim. sans Vert., p. 117); and in 1819 (Hist. Nat. Anim. sans Vert., 
vol. vi, pp. 63-64) T. scabra for fig. 1, T. nodulosa for fig. 2, T. navis 
for fig. 3, and T. aspera for fig. 4; of the latter T. nodulosa Lam. 1819 
is invalid and T. aspera is a synonym of T. nodulosa Lam. 1801. It 
has usually been stated ft that T. nodulosa Lam. 1801 is the first 
specific name to be coupled with that of the genus and that according 
to Opinion 46 (quoted above) this species is zpso facto the type. But 
if it be held that all the unnamed species of Bruguiére 1797 are 
available for subsequent type designation, then it does not follow 
that the naming of one of them in 1801 excludes the other species 
as possible types, since it has been held (Opinions 79 and 81) that 
rigidly construed, Lamarck’s 1801 Syst. Anim. sans Vert. is not to 
be accepted as designation of type species. Stewart (1930, pp. 88-90), 
who has recently gone very fully into the matter, has accepted the 
latter position, but has shown that the first valid designation (on 
the assumption that the genosyntypes are the unnamed species 

* Information published by Stewart (1930, p. 88). 
tT But vide infra. 


Lower Cretaceous Lamellibranchia from Northern Zululand. 319 


figured by Bruguiere 1797) is that of Gray (Proc. Zool. Soc. Lond., 
1847, p. 197) who cited Trigonia as of Lamarck 1801, so that T. 
nodulosa became the type by monotypy on Gray’s interpretation. 

It may well be doubted whether specifically unnamed forms can 
be accepted as genosyntypes, though the Rules and Opinions are not 
clear on this point. Opinion 35 states that “In determining the 
type of a genus, the selection must be confined to species included 
under the generic name in question at the time of its original publica- 
tion, regardless of the fact whether they were named binomially or 
not,” and, further, Article 30 states wnter alia that species must be 
excluded from consideration in determining the types of genera 
“that were not included under the generic name at the time of its 
original publication.”” Nomenclature can know nothing of a species 
unless and until it has a specific name, and where, as in this instance, 
no named species were included under the generic name at the time 
of its original publication we must fall back on the first-named species 
published in connection with the genus. Unnamed figures under a 
generic title cannot be regarded as a proper indication of species, 
though the figures of Bruguiere 1797 serve to clarify the brief 
diagnosis of Bruguiere 1789. The whole spirit of Article 30 and 
Opinion 35 is against accepting a genus proposed without a proper 
indication of species, and Article 30 must have precedence over 
Opinion 46 which would allow the genus (“If it is not evident how 
many or what species are involved, the genus contains all the species 
of the world which would come under the generic description”’). 
The safest course to adopt in the present instance, and the one most 
in accord with the spirit of Article 30, is to quote the genus as of the 
first author to indicate a species, with Bruguiére 1789 in brackets, 
thereby giving recognition to the coining of the generic name by the 
earlier author; assuming for the moment that the earliest use of 
Trigoma with indication of species is Lamarck 1801 (Syst. Anim. 
sans Vert.), the genus should be quoted “ Trigonia (Bruguiére 1789) 
Lamarck 1801, genotype TZ. nodulosa Lamarck 1801, type by 
monotypy.” This point of view has been adopted by Marwick 
(1932, p. 506), who has pointed out that biologists are not unanimous 
in their treatment of genera based on figures alone, without a specific 
name, at the time of publication, and has accordingly suggested 
that the simplest and best course is to treat a genus as not valid 
until it is published along with a specific name that can be identified. 
He is inclined to quote Trigonia as of Lamarck 1801, unless the 
uncertainty expressed by certain authors as to the identity of 


320 Annals of the South African Museum. 


T. nodulosa Lam. 1801 necessitates falling back on the next earliest 
reference, which is Trigonia Lam. 1804 (Ann. Mus. Nat. Hist., vol. iv, 
p. 355) with monotype 7. margaritacea Lam. 1804. 

It would appear, then, that unless an earlier reference than Lamarck 
1801 can be found, the genotype of Trigonia must be T. nodulosa 
Lamarck 1801, whether the genus be quoted as of Bruguiére 1789 
or Lamarck 1801, and in the former case whether the genotype was 
fixed by the naming of a species by Lamarck in 1801 or by the 
subsequent designation by Gray in 1847. This species has indeed 
been very generally regarded as the type, e.g. by Fischer (1887, p. 994) 
and Stewart (1930, p. 89), while Cossmann (1912, p. 62) and Marwick 
(1932, p. 506) have questioned the validity only on the grounds that 
the species was not well established, and Crickmay (1932, p. 446) 
admits that he long regarded the species as “‘monotype.”’ 

It follows from the above argument that Children (Lamarck’s 
Gen. Shells, 1823, p. 48) and Stoliczka (1871, p. 310) were in error in 
accepting the living species T'. margaritacea Lam. 1804 (= T. pectinata 
Lam. 1819) as genotype. This species can only become the genotype 
of Trigonia if both Trigonia Bruguiere and Trigonia Lamarck 1801 
are regarded as not valid, in which case Trigonia Lamarck 1804 would 
have to be accepted, being the next earliest reference. This would 
only arise if T. nodulosa Lam. 1801 were regarded as indeterminable, 
as Marwick (1932, p. 506) has put forward as a possibility, following 
Cossmann (1912). The figure on which T. nodulosa Lam. 1801 was 
based is by no means bad, and more recently the original specimen 
of T. nodulosa Lam. 1819 has been figured by Favre (1914, Cat. Illus. 
Coll. Lam., Mus. d’Hist. Nat. Geneéve, pl. 35, fig. 253,a@, 6); the 
reference is given by Stewart (1930, p. 89), who states that the figures 
do not differ much from the original figure of T. nodulosa Lam. 1801 
and that it seems probable that they represent, if not the same 
species, closely related forms. The species T. nodulosa Lam. 1801 
is a delicately sculptured member of the Clavellatae, and even if 
the lectotype cannot with absolute certainty be identified with 
T. herbert. Bigot or any other species, its affinities can be expressed 
within such narrow limits that there could be no doubt as to the 
meaning of T'rigonia (sensu stricto) by adopting the species as genotype. 
Marwick (1932, p. 506) has pointed out that the selection of an allied 
species, T. clavellata Sowerby, as neogenotype by Cossmann (1912, 
p. 62) is quite invalid, and the same opinion is presumably held by 
Stewart and Crickmay. 

We are indebted to Crickmay (1932, p. 446) for bringing to light 


Lower Cretaceous Lamellibranchia from Northern Zululand. 321 


an earlier reference than Lamarck 1801, a reference that has appar- 
ently been entirely overlooked by previous investigators, and which 
has necessitated a re-examination of the whole problem. In his 
Prodome d’une nouvelle classification des Coquilles (Mém. Soc. Hist. 
nat. Paris, 1799), on pp. 86, 87, Lamarck gives a short diagnosis of 
Trigoma, followed by two references, one to certain named species 
figured by Hermann in 1781. The diagnosis reads as follows: 
“Trigonie. Trigonia. Coq. inéquilaterale, subtrigone; charniére & 
deux grosses dents plates, divergentes, et sillonnées transversale- 
ment,” and is followed by “Encycl. t. 237; Naturforsch, 15e livraison, 
t.iv.” The first reference is of course to the first plate of Bruguiére 
1797, and the second has been identified by Crickmay with a plate 
accompanying a paper by Hermann (Brief iiber einige Petrefacten, 
Der Naturforscher, Bd. 15, t. iv, Hall, 1781). On that plate appear 
16 figures of several species of Lamellibranchs bearing valid specific 
names, all Trigoniae except figure 1, which appears to be a Phola- 
domya. The Trigonae (figs. 2-16) are grouped in four species under 
the generic title “Venus oder Donaz,”’ viz. sulcata, tuberculata, 
nodosa, and dubia.* This would appear to be the earliest reference 
of named species to the genus Trigonia, and T. nodulosa Lam. 1801 
would only be the genotype if we accept the contention that the 
genosyntypes must be chosen from among the unnamed figures of 
Bruguiere 1797. Two courses are open on the available data—(1) 
to accept Bruguiére 1797 as fixing the genosyntypes, and (2) to accept 
Lamarck 1799 as fixing the genosyntypes on the ground that it 
contains the earliest reference to a species by name; the first course 
would lead to accepting as genotype a Clavellate Trigonia, as out- 
lined above; the second course would lead to accepting as genotype 
a Costate Trigonia, T. sulcata (Hermann), type by subsequent 
designation (Crickmay, 1932). The ambiguities and contradictions 
in the International Code make it difficult to come to a decision on 
this point, but it would seem, from the arguments set forth above, 
that the second course is preferable, and the writer accordingly 


* T am indebted to L. R. Cox for looking up the references Lamarck 1799 and 
Hermann 1781 at the British Museum. The figures on Hermann’s pl. iv are 
named as follows :— 

Fig. 1. Cardium tortuosum. 

Figs. 2-4, 9, 10. Venus oder Donax sulcata. 

Figs. 5, 8. Venus oder Donax tuberculata. 

Figs. 6, 7, 15, 16. Venus oder Donax nodosa. 

Figs. 11, 12. Venus oder Donax dubia. 

Figs. 13, 14. Venus oder Donax zwischen sulcata und dubia. 


322 Annals of the South African Museum. 


accepts T. sulcata (Hermann) as genotype of Trigonia (Bruguiere 
1789) Lamarck 1799. The original figures of the lectotype of this 
species, a fine representative of the Costatae, have been reproduced 
by Crickmay (1932, pl. 1, figs. 1, 2); these correspond to Hermann’s 
figures 4 and 3 respectively, two views of the same shell. 

The writer is not, however, in agreement with the arguments set 
forth at some length by Crickmay (1932, pp. 447-451) in favour 
of accepting Hermann’s species as genosyntypes, even though the 
arguments lead to the same result. A fairly clear case seems to have 
been made out above for accepting these species by reason of priority 
of reference to the genus, assuming that unnamed figures are not 
admissible. Crickmay raises two objections against the customary 
acceptance of T. nodulosa Lamarck 1801 (or of T. margaritacea) as 
genotype. The first objection, that the species do not conform 
strictly to the original generic diagnosis in that they have ovate 
rather than triangular shells, can scarcely be taken seriously, depend- 
ing as it does on the use of the word “‘triangular”’ in this connection. 
The older authors apply the terms triangular and trigonal to the 
majority of the Trigoniae, and even if the term is used in a more 
exact sense it can be used for a considerable range of species in the 
genus, including many representatives of the Clavellatae, as can 
readily be seen, e.g. by inspecting the plates of Lycett’s monograph. 
Indeed, in many species of Clavellatae the shape is as markedly 
triangular as in the majority of the Costatae, and in several cases 
paired species from the two groups can scarcely be distinguished 
in outline. There are no grounds whatever for the statement that 
only the Costatae have triangular shells, and T. nodulosa might well 
have been described as triangular by many authors. It does not 
follow, therefore, that it is probable that Bruguiére had the Costatae 
particularly in mind when founding the genus, and that therefore 
this group ought to be regarded as typical. 

The second objection was raised against the custom permitted by 
Opinion 46 of the International Code “of genotype selection literally 
from among all the species of the world,’ in the case of a genus pub- 
lished without species. The restriction suggested by him is novel, 
but can scarcely be maintained. “It is impossible for a species un- 
known to the author of the genus to be in any real sense the genotype. 
Only species named by the time the genus name appeared can possibly 
have served an author for his conception of his genus. Such previous 
species are the only concrete things on the basis of which a genus 
published without species can be known in binomial nomenclature 


Lower Cretaceous Lamellibranchia from Northern Zululand. 323 


until species are joined to it. It ought to make no difference that 
these previous species are not coupled with the genus name until 
later; they may be correlated with it by their agreement with the 
generic diagnosis. Among them only a genotype ought to be sought.” 
He then proceeds to show that the species of “Venus oder Donaz,”’ 
published by Hermann in 1781, fall under the diagnosis of Trigonia 
published by Bruguiére in 1789, and accepting them as being among 
the possible genosyntypes, designates one of them as genotype. The 
argument seems to be based on the assumption that the genotype 
must be a species known to the author of the genus, but one might 
ask what guarantee there is that Bruguiére was aware of or had 
particularly in mind Hermann’s species when he founded Trigonia; 
it is indeed conceivable that Bruguiére had in mind only certain 
specimens in the collections accessible to him, of unpublished species, 
and was unaware of the fact that certain previously published species 
would fall under his generic diagnosis. The essential point in limit- 
ing the choice of genotypes is the fact of reference of species by the 
author at the time of original publication of the generic name; 
Article 30 and Opinion 35 are quite clear on this point. Article 25 a 
and Opinion 46 apparently allow a genus to be treated as valid when 
it is proposed without species, but it may be doubted whether any 
real knowledge of the genus can be obtained until a subsequent 
author associates named species with the generic name, and if that 
is so the subsequent author should be regarded as the real author 
of the genus. 

Crickmay’s proposal brings in another difficulty which was not 
considered by him; if, in a case of this kind, the genotype is among 
the species described previously under other generic names, the usual 
priority rules should apply, and the genotype would have to be the 
earliest or at least among the earliest of such species. It may be 
doubted whether the species of Hermann 1781 are the earliest species 
of what we now call Trigonia; there may be several earlier species 
which have not been recognised for what they are, either because 
of the rarity of the publication in which they appear or because of 
some doubt as to their agreement with the original generic diagnosis. 
It would be possible for an author to unearth, successively, older 
species, some possibly of very doubtful affinities, which in his opinion 
agree with the original generic diagnosis, and to designate each in 
turn as genotype—surely a reductio ad absurdum ! 

Crickmay (1932, p. 449) has sought to strengthen his conclusions 
by stating that the species of Hermann 1781 were reproduced in 


324 Annals of the South African Museum. 


Bruguiére’s plates of 1797. The writer is indebted to Mr. L. R. Cox 
for the statement that Bruguiére’s figures are absolutely different from 
those of Harmann, and that there can be no question of their being 
copies. The identification of figures on Bruguiére’s plates with 
Hermann’s species is merely a statement of opinion, and it would 
seem from a comparison of the figures reproduced by him that he is 
probably incorrect in some instances. For example, on pl. xi, figures 
1 and 2 are said to be copies of Bruguiére; pl. cexxxvii, figs. 1 a, 
1b, which on p. 449, footnote, are said to be the equivalent of Her- 
mann’s sulcata; it may be doubted whether these figures represent 
the same species as the figures of the lectotype of T. sulcata repro- 
duced on Plate I. 

Synonyms of Trigonia sensu stricto.—In 1823 Sowerby (Min. Conch., 
vol. v, p. 40) substituted the name Lyridon (emended Lyriodon 
Bronn) for Trigonia on the grounds that Trigonia had previously 
been used for a plant. Since this does not constitute a case of pre- 
occupation under the existing rules, Lyridon is a strict synonym 
of Trigonia, as has been pointed out by Cossmann (1912, p. 62) and 
Stewart (1930, pp. 88-90). Lyridon, and its emendations Lyriodon 
and Liriodon are therefore not available for use for the Costatae as 
has been done by Rollier (Mém. Soc. Pal. Suisse, 1912, vol. xxxvii, 
p- 65), Deecke (1925, p. 68), Dietrich (1933, p. 32), and others. 

Stewart (loc. cit.) has pointed out that Lyrodon Goldfuss is virtually 
in the same category as these other emendations, but might be 
revived, as Goldfuss did not actually say he was renaming Trigonia. 
Stewart accordingly designated as type the Clavellate T. aspera 
Lamarck, which is synonymous with T. nodulosa Lam. 1801, in 
order to make Lyrodon Goldfuss an objective synonym of Trigonia; 
but if 7. sulcata (Hermann) is accepted as genotype of Trigonza (follow- 
ing Crickmay 1932), then Lyrodon might possibly be revived for the 
Clavellatae, for it would have priority over Myophorella Bayle. But 
Goldfuss certainly used the term as an objective synonym of Trigonza 
even if he did not explicitly state so, and was clearly following 
Sowerby and Bronn, hence the term must be a strict synonym of 
Trigona. 

Myophorella Bayle 1878 (Explic. Carte Géol. France, Foss. Prine. 
des Terr., vol. iv, pl. cxx) was based on two figured species, M. nodu- 
losa (Lam.) and M. muricata (Roemer), both Clavellatae. The genus 
was stated to be a synonym of Trigona by Cossmann (1912, p. 63) 
ex eodem typo, which seems to imply the designation of M. nodulosa 
(Lam.) Bayle as genotype. If T. nodulosa Lam. 1801 is genotype 


Lower Cretaceous Lamellibranchia from Northern Zululand. 325 


of Trigonia then Myophorella becomes a synonym even if there is 
doubt as to the identity of the type species. On the other hand, 
with 7. sulcata as genotype of Trigonia (Costatae), Myophorella is 
available for the Clavellatae with 7. nodulosa (Bayle) non Lamarck 
as type (Crickmay, 1932, p. 458). 

Classification of the Trigonias.—Until comparatively recently there 
has been little departure from the position taken up by Agassiz as 
long ago as 1840 as regards the classification of the Trigoniae, and 
it has been customary to refer the Trigoniae to one huge genus with- 
out sub-division on a Linnaean basis. Agassiz (1840, pp. 7-10) 
recognised eight groups of species, to which French names were given 
and whose “types” were in most cases designated, and these groups 
were believed to be more or less natural. Lycett (1872-1879) added 
a ninth group and translated the group-names into Latin plurals, 
while subsequent authors have admitted additional groups, by sub- 
division or otherwise, but without seriously modifying the original 
scheme. 

The necessity for smaller units has been pointed out by several 
recent workers, but it is interesting to compare the views of Lycett 
and others on this matter. Lycett (sbid., p. 217) had been tempted 
to sub-divide the genus into a number of distinct genera, but the 
results of his observations led to the perception of a general resem- 
blance between the several groups of species in features of sufficient 
importance to induce him to regard them as forming part of one great 
whole. Cossmann (1912, pp. 81-84) in his essay on the evolution 
of the Trigoniae was impressed by an essential unity among the 
Mesozoic members of the genus, and after pointing out the important 
structural differences between these and the post-Cretaceous Austral- 
asian species (Agassiz’s Pectinatae, Cossmann’s genera Hotrigonia 
and Neotrigonia), was led to remark “‘tousles . . . groupes présentent 
des transitions graduelles d’un groupe 4 Vautre, ce qui démontre 
Vinanité du systéme adopté par Agassiz” (ibed., p. 62). 

On the other hand, few would agree that the accepted groups are 
as artificial and indistinctly limited as Cossmann would have us 
believe, and several recent workers have felt that the time is ripe 
for sub-division. The matter has been discussed at some length by 
Crickmay (1932, pp. 443-445), who has emphasised the discrepancy 
between the comprehensiveness of Trigonia and the fine distinctions 
drawn between genera in the Ammonoidea and other groups, living 
as well as fossil. COrickmay has also drawn attention to the fact 
that in the Trigoniae the evolution of forms was neither so rapid nor 


| oe: 


326 Annals of the South African Museum. 


so adventitious as to obscure the course of phylogeny. The retention 
of a distinctive juvenile ornamentation in many Trigoniae has indeed 
been of very great value in the determination of relationships and 
discrimination of natural groups, and has been successfully utilised 
by Kitchin (1903) and others. Classification based on adult char- 
acters alone has certainly led in the past to the proposal of certain 
groups of a very artificial character, the phenomenon of homeo- 
morphy, whose prevalence has been amply demonstrated by modern 
palaeontological research in many groups of animals, being well dis- 
played in the Trigoniae. It would appear, however, that there are 
grounds for believing that a number of the narrowly limited genera 
or sub-genera that have been proposed during the last ten years are 
more or less natural groups, though homeomorphy is sometimes 
evident, e.g. the distinction between 7. vau and T. v-scripta pointed 
out by Kitchin (1903, p. 65; 1908, p. 112) based on the character 
of the juvenile concentric ribbing (see remarks below under Lotrigonia). 
It is evident that an adequate and full conception of the evolution 
of the Trigoniae will only be obtained after a very extensive study 
of the vertical and horizontal distribution of the Trigoniae, and the 
present tendency to recognise a number of genera or sub-genera 
cannot but materially assist in the elucidation of the problem. 
Deecke, Crickmay, Marwick, and others have proposed the new 
divisions with the status of genera. To a great extent the distinction 
between genera and sub-genera is purely arbitrary and has admittedly 
no legal sanction, but nevertheless the writer favours the use of these 
new divisions as sub-genera, following the lead given by Dietrich 
(1933). One cannot but be impressed by the extraordinary constancy 
of the characters of the dental apparatus of the Trigoniae throughout 
the Jurassic and Cretaceous periods. Alien elements of pre-Jurassic 
and post-Cretaceous age have rightly been separated as genera, but 
compared with these the Jurassic and Cretaceous Trigoniae form in 
this respect a unit, if an unwieldy one. The variations in dental 
characters are confined to differences in the relative sizes of teeth, 
differences in the robustness of the teeth, differences in the degree 
of bifidity shown by the left central tooth, and differences in the 
degree of divergence shown by the teeth, features which in the main 
are closely linked up with differences of external form. As a conse- 
quence genera have been proposed largely, if not entirely, on a con- 
sideration of external form and ornamentation, and linkage between 
recognised groups is frequently indicated by the character of the 
ornament in the youthful stage. The late Dr. F. A. Bather in his 


Lower Cretaceous Lamellibranchia from Northern Zululand. 327 


Anniversary Address to the Geological Society of London on 
“Biological Classification: Past and Future” (Q.J.G.S., Proc., 
vol. lxxxin, 1927, pp. [xxxix—xc) emphasised that classification has 
a practical as well as a theoretical aim, and that the discrimination 
of units must not be confused with the practical aim of assortment 
of units for convenience of reference. The fact of seriation, which 
is obscured by the multiplication of genera, is more important than 
the discontinuities which the multiplication of genera is designed to 
illuminate. It appears to the writer that the acceptance of the new 
groups of Trigonia as sub-genera is to be commended both on prac- 
tical grounds and for the emphasis it gives to a certain uniformity 
of structure, however much that uniformity may be disturbed by 
differences in sculpture. As genera, possibly grouped together as 
units of higher rank, that fundamental uniformity is no longer 
reflected in the nomenclature. It may be doubted whether the 
elevation of the sections of Agassiz and others, virtually sub-genera, 
into sub-families (van Hoepen, 1929) is warranted on the evidence. 

One cannot leave this subject without expressing regret at the 
tendency, as shown in the choice of generic names and in the desig- 
nation of genotypes, to ignore the names and types of the groups 
whose recognition is very largely the result of the patient labours 
of workers during the nineteenth century. It appears to the writer 
that it would only be courteous to earlier workers to perpetuate as 
far as possible the names given by them, as has been done by Deecke. 
Since the “types” designated by Agassiz and others are in the main 
well-known species, some of which have been accepted as “‘types”’ 
for the greater part of a century, they should wherever possible be 
designated types of the new genera or sub-genera. It is therefore 
to be regretted that van Hoepen (1929) has thought fit to base 
certain genera on comparatively little-known species from a remote 
locality, species that do not differ in features of more than specific 
value from the older “types” of Agassiz, Kitchin, and others, that 
have been amply described in various publications. 

The Generic Names proposed by Deecke.—The writer cannot too 
strongly deplore the casual way in which Deecke (1925, p. 68) has 
proposed certain new generic terms for groups of Trigoniae, but at 
the same time it would appear that these highly suitable names must 
be regarded as validly proposed. 

Deecke introduced the terms Laevitrigonia, Scaphotrigonia, and 
Scabrotrigonia for the Laeves or Glabrae, Scaphoideae, and Scabrae 
respectively, thus perpetuating the groups recognised by Agassiz 


328 Annals of the South African Museum. 


and Lycett, but failed to give diagnoses or to designate genotypes. 
Van Hoepen (1929, p. 3) held these names to be nomina nuda and 
has been followed “‘in the interest of uniformity” by Stewart (1930, 
p. 90), as well as by Crickmay (1932, p. 452). 

There cannot be the slightest doubt as to Deecke’s intention when 
publishing these names. After referring to the groups of Agassiz by 
name, Deecke goes on to state: “Steinmann meinte, die alten 
Agassizschen Gruppen beséssen den Wert von Untergattungen und 
hat damit wohl fiir die Laeves, Costatae und vielleicht fiir die 
Scaphoiden recht. Fiir die Costaten fiihrte Rollier den alten Sowerby- 
Bronnschen Namen Lyriodon als Untergattungsnamen ein und fiir 
die beiden anderen waren Laevitrigonia und Scaphotrigonia durchaus 
brauchbar. Ihnen lhesse sich noch Scabrotrigonia anschliessen.” 
The wording of the last two sentences is unfortunate and Crickmay 
has objected inter alia that the names were not formally proposed, 
but the names appear in print and could be interpreted as genera, 
and must be accepted if there is sufficient indication as to the author’s 
intentions. The names are apparently, though not expressedly, 
tautonyms of Lycett’s Latin plurals of three of the group-names 
proposed by Agassiz. Crickmay implied that the names are accom- 
panied by no indication and are therefore nomina nuda in spite of 
the tautonymy. But there cannot be any doubt whatever, from 
the brief discussion which precedes the proposal, that Deecke had 
in mind certain identifiable species-groups when proposing the genera, 
or that Laevitrigonia, Scaphotrigonia, and Scabrotrigonia were in- 
tended for the Glabrae, Scaphoideae, and Scabrae of Agassiz re- 
spectively. There is a sufficient indication here of the species-groups 
to which it was intended that these names should apply, and from 
Article 25 * and Opinion 1+ of the International Rules it would 
appear that the names are valid. 

The objection raised by van Hoepen and also by Crickmay that 
the species-groups concerned are heterogeneous does not affect the 
validity of the proposal, since the genera can be restricted by the 
subsequent designation of genotypes. 

It is interesting to find that Stewart (1930, p. 89) was compelled 
to admit that there was little doubt as to what these names were 
proposed for, and had accepted the names and designated types for 
them in manuscript. His abandonment of the names “‘in the interest 
of uniformity,” following van Hoepen’s rejection, has little to com- 
mend it; the rules of nomenclature should be strictly adhered to. 


* TX° Congrés Internat. Zool. 4 Monaco (Rennes, 1914), p. 901. T Jbid., p. 909. 


Lower Cretaceous Lamellibranchia from Northern Zululand. 329 


Deecke’s genera have been accepted by Dietrich (1933, p- 35), and 
Mr. L. R. Cox, M.A., F.G.S., in a letter to the writer, has expressed 
the opinion that the genera were validly proposed. 

It had been the present writer’s intention to designate genotypes 
for Deecke’s three genera,* choosing the “types” of the corresponding 
groups recognised by earlier workers. Dietrich (loc. cit.) has rightly 
designated 7. gibbosa Sow. type of Laevitrigonia, this being the 
earliest designation that the writer has been able to find. The 
writer hereby designates 7. navis Lamarck genotype of Scapho- 
trigonia, and T. scabra Lamarck genotype of Scabrotrigonia, in the 
belief that designations have not hitherto been made. 

The Generic Names Proposed by van Hoepen.—The rather numerous 
species of Trigonia in the Cretaceous deposits of Zululand in South 
Atrica have been studied by van Hoepen (1929). In his paper 
twenty-one species and varieties of Trigonia from the Umsinene 
River in Zululand and from Pondoland are arranged under no less 
than nine genera and four sub-families, while a tenth genus was 
founded on a species from the Uitenhage Series farther south. The 
minuteness of some of these sub-divisions may be gauged from the 
fact that the Scabrae, even after the elimination of the doubtful 
group Megatrigoma (known to others as the group of T. conocardii- 
formis), are represented in South Africa by six genera, and that the 
lengthy diagnoses of three of these genera are so closely similar as 
to be virtually identical. The author claims to have made an 
attempt to classify the Zululand species on a phylogenetic basis 
(ibid., p. 3), but a perusal of his paper fails to reveal much of phylo- 
genetic significance; the species are well described, but throughout 
the paper there is a fine disregard for the work of previous investi- 
gators, and the author makes no attempt whatever to trace the phylo- 
geny of the groups he distinguishes. Van Hoepen appears to have 
solved the problem of classification of some twenty species of Trigonia 
by designating half of them genotypes of new genera. Some authors 
may find it a convenient solution of their taxonomic problems to 
consider only a small number of local species and make most of them 
types of distinct genera, but some of the genera thus formed are not 
likely to have any systematic value when applied elsewhere. Van 
Hoepen is mistaken if he assumes that the mere multiplication of 
genera, and consequent increase of a somewhat burdensome nomen- 


* The writer had prepared a discussion on the Trigoniae in 1930 for a paper 
on the Cretaceous of Angola, in which designations were made. The paper has 
unfortunately not yet been published. 


330 Annals of the South African Museum. 


clature, has any essential bearing on the subject of phylogenetic 
relationships. 

The novelty of outlook displayed by van Hoepen’s work may be 
judged from the following examples: (1) Kitchin, an experienced 
worker and acknowledged authority, was scarcely able to separate 
from T. ventricosa (Krauss) (Uitenhage species) certain specimens 
from the Oomia Beds of Cutch after a careful examination of numerous 
specumens from both areas (Kitchin, 1908, p. 95)—yet van Hoepen, 
who presumably was relying only on Kitchin’s figures and descrip- 
tions of the Oomia specimens and not on actual examples, not only 
gave the latter the new name Pisotrigonia parva, but made the Uiten- 
hage species the genotype of a new and different genus ! (van Hoepen, 
p. 22); (2) In the Umzamba Beds in Pondoland there is a very 
abundant small Trigonia long known under the name of T. elegans 
Baily; van Hoepen has not only split this species into three distinct 
species (a step which an examination of some hundreds of finely 
preserved topotypes shows to be scarcely warranted), but has followed 
the curious course of designating as genotype of Linotrigonia, not 
the well-known 7. elegans with which he was familiar, but his new 
species L. linifera, founded on a fragmentary mould of a left valve and 
said to differ from 7’. elegans chiefly in its larger size (vbid., p. 16). 

Some of the genera proposed by van Hoepen will prove useful in 
future work, but it would appear that his fine sub-division of a portion 
of the older group of the Scabrae can scarcely be maintained. The 
following are the groups whose Zululand representatives were 
investigated by van Hoepen:— 

(1) Megatrigoma (type M. obesa van H.) was proposed for a single 
species that was compared only with the elongate Scabrid T. rogersi 
Kitchin. The sub-genus is equivalent to the previously recognised 
group of T. conocardiuformis, the close resemblance of the species 
concerned having been overlooked by van Hoepen. 

(2) Iotrigonia (type I. crassitesta van H.) should prove useful for 
the convergent groups of T. v-scripta and T. vau distinguished by 
Kitchin. Van Hoepen, in including both of the latter species in 
Totrigonia, fails to mention the distinction drawn by Kitchin, and 
it is not clear as to which group the type species belongs. 

(3) The type species of the six genera of Scabrae proposed by van 
Hoepen are from the Cretaceous of Zululand, the Senonian of Pondo- 
land, and the Neocomian of the Uitenhage basin, and the majority 
would have been regarded as typical Scabrae by other workers. It 
is doubtful whether the distinctions made between some of these 


Lower Cretaceous Lamellibranchia from Northern Zululand. 331 


genera are of much significance, and one or more of them would be 
included in the synonymy of Scabrotrigonia Deecke. The genera 
concerned are Péerotrigonia, Acanthotrigonia, Linotrigonia, Piso- 
trigoma, Ptilotrigonia, and Rinetrigonia. Crickmay (1932, pp. 460- 
463), in giving brief diagnoses of these genera, has referred to the low 
status of some of them. 

(4) Rutetrigoma (type R. peregrina van H.) was proposed for a 
single species from South Africa which was compared with certain 
Kuropean species which have hitherto been included in Bigot’s 
group of the Hxcentricae. The group-names Glabrae and Excen- 
tricae were not mentioned, but it would appear that R. peregrina is 
a fairly characteristic member of the latter group, and Rutitrigonia 
can therefore be used in place of the group name Excentricae. The 
substitution of the “type” of the latter by a later species from a 
remote locality must be deplored. 

(5) Pleurotrigoma (type T. blanckenhorn R. B. Newton) is founded 
on a single isolated species of Costate ancestry and appears to be 
worthy of retention as a sub-genus. 

Names Given to Extra-South African Species by van Hoepen.—Two 
new specific names were proposed by van Hoepen for species from the 
northern hemisphere, and as there is some danger that these validly 
proposed names may be overlooked, reference is made to them here. 

Ptilotrigonia crassicostata van Hoepen (1929, p. 26) was proposed 
for Trigoma aliformis dOrbigny (Pal. Frangaise, Terr. crétacés, 
vol. 11, 1843, pl. 291, figs. 1-3), following the remark of Lycett that 
the latter was distinct from 7. aliformis Park. 

Pisotrigona parva van Hoepen (1929, p. 22) was proposed for the 
Oomia shells previously regarded by Kitchin as scarcely distinguish- 
able from the South African TZ. ventricosa (Krauss). P. parva 
(Kitchin, 1903, p. 104, pl. x, fig. 4—the other figures were not indi- 
cated by van Hoepen) must be very close to 7. ventricosa, since Kitchin 
(1908, p. 95) states that the points of difference do not suffice even 
for the satisfactory recognition of two well-defined local races.* 


Sub-genus Megatrigonia van Hoepen 1929. 


The sub-genus Megatrigonia was proposed by van Hoepen (1929, 
p. 3), with the rank of a genus, united with Jotrigonia in the 
sub-family Megatrigoninae, for the single species M. obesa van 

* Cox (Pal. Indica, 1935) says of the Indian shells that “their ornamentation 


is identical with that of South African specimens, and there is no justification for 
referring the Indian form to a distinct species . . . far less to a distinct genus.” 


332 Annals of the South African Museum. 


Hoepen from the Lower Cretaceous of Zululand. The type species 
was compared only with the superficially similar Uitenhage species 
T. rogersi Kitchin, regarded by Kitchin (1908, p. 101) as a normal if 
elongated representative of the section Scabrae and placed in another 
sub-family by van Hoepen himself. M. obesa and T. rogersi can 
only be remotely related. Van Hoepen overlooked the very striking 
similarity indicative of close relationship between his M. obesa and 
the well-known and rather isolated Uitenhage species T. conocardw- 
formis (Krauss) (Kitchin, 1908, p. 119, pl. vu, figs. 2-4), and the 
still greater resemblance to the shells from Hast Africa identified 
with the Uitenhage species by Lange (1914, p. 235, pl. xix, fig. 1). 
The omission is curious, since van Hoepen was presumably familiar 
with Kitchin’s paper, in which the peculiarities of 7. conocarduformis 
are finely described, and which includes excellent figures showing 
area and escutcheon; the adult shell, however, has not been adequately 
figured up to the present. The latter species agrees with M. obesa 
in all essential features, and differs only in details such as suffice to 
distinguish closely allied species; the average field geologist acquainted 
with the Uitenhage fauna would scarcely have hesitated to apply the 
Uitenhage name to the Zululand shells. 

The sub-generic diagnosis has been given by van Hoepen and sum- 
marised by Crickmay (1932, p. 460), and may be stated as follows:— 
Shell very large, considerably longer than high, highest in the 
anterior third, the posterior part narrower and produced, roughly 
pear-shaped in outline. Anterior margin very broadly convex and 
passing gradually into the convex ventral margin; postero-dorsally 
concave, the margin nearly straight or concave and rather long; 
posterior margin short. Umbonal region prominent. Moderately 
inflated in the anterior half and very gradually becoming flattened 
towards the posterior. Area smooth and rounded, except in the 
early neanic stage, and becoming progressively wider when traced 
to the posterior margin. Hscutcheon smooth, lanceolate and deeply 
concave. Dorsal carinae and longitudinal costellae absent, the 
former, however, represented by weak folds in the neanic stage. 
Ornamented on the flank with numerous rounded, oblique costae; 
the anterior costae massive, smooth to more or less nodose, curving 
downwards and then forwards; the costae becoming progressively 
finer towards the posterior, tending to be directed vertically and to 
be less nodose. Umbo concentrically ribbed in the early neanic 
stage, the ribs passing on to the area. Dentition strong, the left 
central tooth very markedly bifid. 


Lower Cretaceous Lamellibranchia from Northern Zululand. 333 


The sub-genus occupies a somewhat isolated position among the 
Trigomae, and is well worthy of recognition. Lycett (1879, p. 210) 
came to the conclusion that the species T. conocardiiformis was un- 
doubtedly associated with the Scabrae, but Kitchin (1908, p. 123) 
stated that definite indications of such a relationship are not to be 
recognised. The group occurs in Neocomian strata from Tanganyika 
to South America. 


Trigoma (Megatrigonia) obesa (van Hoepen). 
(Plate XX XVIII, figs. 1-3; Plate XX XIX, figs. 1, 2.) 


1929. Megatrigonia obesa van Hoepen: Krytfauna Soeloeland, pp. 
4-6, pl. i, figs. 1, 2; pl. 11, figs. 1-3. | 

Material.—The type set was collected by van Hoepen from his 
Ndabana Beds on the Umsinene River, Zululand. In the collections 
from the Pongola area there are about 30 specimens. Mr. H. F. 
Frommurze and Mr. J. 8. Hutt collected 10 specimens along the 
Mfongosi tributary of the Pongola River (T.M. 1763, 1778, 
(Slee o2 SIS. 19095 SAME 10784, 10785). Dri. S. ~ H: 
Haughton and Dr. A. W. Rogers collected about 20 specimens at 
the same locality, distributed in the zones as follows, the numbers 
in brackets referring to the number of specimens found: ZA (2), 
Z2 (1, juvenile), Z3 (7), Z4 (4), Z5 (1), Z8 (5), Z12 (1). A single 
specimen was collected farther south on the Myesa stream, MI (1.) 

The material includes a number of weathered specimens, but some 
are fairly well preserved and complete. I have not seen the details 
of the dorsal area, escutcheon, and dentition, which have been well 
described by van Hoepen. 

Description of the Species.—The following is based as far as possible 
on the new collection from the Pongola, but is in agreement with that 
given by van Hoepen; a free translation of portions of van Hoepen’s 
paper is included so as to complete the description. 

“Shell very large, considerably longer than high, highest in the 
anterior third, the posterior part narrower and produced, roughly 
pear-shaped in outline. Anterior margin very broadly convex, and 
passing quite gradually into the less convex ventral margin; the 
anterior margin with its greatest forward convexity a little below 


* Where quotations from van Hoepen are given under the descriptions of the 
species of Trigonia, it is to be understood that the writer has given an English 
translation. 

VOL. XXXI, PART 3. 28 


i 


334 Annals of the South African Museum. 


the middle of the shell; dorsal part of the anterior margin moderately 
convex to relatively straight and sloping forwards and downwards 
at a moderately steep angle from the umbo. Ventral margin long, 
moderately convex in the anterior half, becoming straighter in the 
posterior half and sloping gradually upwards to meet the posterior 
margin. Postero-dorsal margin moderately long, gently concave, 
sloping very gradually towards the posterior. Posterior margin 
relatively short, less than half the height of the shell, sloping down- 
wards and backwards at a moderately steep angle, meeting the 
postero-dorsal margin in a rounded, very obtuse angle, and meeting 
the ventral margin in a rounded angle of about 90°. Umbonal region 
prominent, the umbo situated between a fourth and a third of the 
length of the valve from the anterior, strongly incurved and curved a 
little posteriorly. Moderately to fairly strongly and evenly inflated 
in the anterior half, very gradually becoming flattened posteriorly, 
with the greatest slope towards the anterior margin, but not anteriorly 
truncate. 

“There is no marginal carina in the adult. In the neanic stage 
the marginal carina is represented by a weak fold, which passes 
posteriorly into a convexity of the valve which becomes progressively 
wider and more indistinct. At the posterior end of the adult shell 
the fold is lost in the uniform convexity of the valve. The inner 
carina is similarly but a weak fold, which becomes indistinct towards 
the posterior and ends near the upper end of the posterior margin. 
Between the carinae and nearer the inner carina, from which it 
diverges a little towards the posterior, runs a shallow longitudinal 
groove which persists to the posterior margin.” 

The juvenile ornamentation of the flank passes into that of the 
adult by gradual increase in the amount of anterior divergence of the 
costae. In the adult the flank is ornamented with numerous rounded 
costae, the whole series of costae appearing to radiate from a point 
just above and behind the umbonal region. On the more inflated 
anterior half of the shell the costae are relatively strong and widely 
separated; towards the posterior the costae quite gradually weaken 
and the interspaces become narrower. The anterior costae are 
strong, rounded, and with broad concave interspaces 2-3 times as 
wide as themselves; they slope steeply downwards and forwards in 
the antero-ventral direction, all more or less convex towards the 
posterior; in front the costae bend gradually forwards so as to 
approach the anterior margin at an acute angle, but they may (as 
in the holotype) bend forwards sufficiently to be roughly perpendicular 


Lower Cretaceous Lamellibranchia from Northern Zululand. 335 


to the anterior margin; those costae which approach the anterior 
part of the ventral margin may bend forwards rather rapidly before 
reaching the margin, even to the extent of diminishing the width 
of the interspaces. In old shells the anterior costae fail to reach 
the anterior and antero-ventral margins, which are then bordered by 
a band marked only by growth lines; the stage at which the costae 
cease varies a little from specimen to specimen, some comparatively 
large shells having little sign of the marginal band or of the relatively 
abrupt forward bending of the costae near the ventral border. In 
the mid-region of the valve the costae are weaker, rounded, separated 
by rounded interspaces about as wide as themselves, and are directed 
approximately vertically, though convex towards the posterior and 
tending to bend forwards before reaching the ventral border. Pos- 
teriorly the costae are progressively weaker and closer together, and 
are directed steeply downwards towards the posterior. 

The flank is ornamented throughout with growth lines, which cross 
both costae and interspaces. A certain periodicity in the growth 
gives rise to a series of quite distinct concentric furrows, commonly 
6-8 mm. apart in the middle part of the anterior region of the shell, 
but closer together towards the margin; these have the effect of 
constricting the costae at more or less regular intervals, so that the 
costae are lower and narrower at the crossing of the concentric 
furrows and raised and swollen between. In some specimens the 
coarsely beaded character of the anterior costae is much in evidence, 
while in others (as in the holotype) it is much less marked; the 
tubercles thus formed may be rounded, oval, or even elongate. 
Towards the posterior the tuberculation is progressively less notice- 
able, and the anterior costae gradually lose. the tubercles and become 
narrower as they are traced towards the area. 

“In the young shell the ornamentation of the flank consists of 
sharp concentric ribs, situated close to one another, which begin at 
the marginal carina, are concave downwards for a very short distance, 
and rapidly pass into a broad downwardly directed curve which 
ends at the anterior margin. The ribs are more or less parallel to 
the ventral margin, but not quite so, one specimen showing that 
the anterior ends of the ribs make an angle of 6° with the growth 
lines at under 5 mm. from the umbo.” At a slightly later stage, 
when the initial pseudo-concentric ribs pass into anteriorly diverging 
costae and gradually increase in thickness away from the area, “‘a 
short initial part of the ribs at the marginal carina remains thin. 
Owing to the bending down of the ribs to continue as very oblique 


336 Annals of the South African Museum. 


costae across the flank, the concavity or embayment of the ribs just 
in front of the carina passes into a sharp bend. This bend does not 
appear in the later costae, which are added successively on the posterior 
side as growth proceeds.” The posterior costae begin below the 
margin of the area. 

“The ornamentation of the area is poor. In the neanic stage the 
area is crossed by costellae, which are parallel to the growth lines 
over the marginal carina, and continue over the area and the inner 
carina towards the umbo. These costellae disappear, and are re- 
placed by very fine growth lines over the area and escutcheon, before 
the stage 1s reached at which the costae lose the sharp bend just in 
front of the marginal carina. The escutcheon is broad and long, and 
for the most part concave.” The area is devoid of costellae after 
the early neanic stage and ornamented only with growth lines. 

The rather strong dentition, characterised by the markedly bifid 
left central tooth with the two forks of the tooth diverging at about 
90°, has been described in detail by van Hoepen and need not be 
here repeated. 

Remarks.—There is a fair amount of variation in the character 
of the ribbing in the Pongola specimens. I have individuals before 
me equally well preserved which differ from one another in the develop- 
ment of the tuberculate character of the anterior costae, varying 
from shells with coarsely beaded costae to shells with rounded costae 
slightly constricted at more or less regular intervals. In the holo- 
type the only adult shell whose external features are shown in van 
Hoepen’s figures, the costae appear to be of the latter type, though 
the original description of the species refers to “the impression of 
tubercles,’ and describes their formation. 

The rapid bending forward of the costae near the ventral margin 
is not characteristic of all the specimens before me, and in no case 
do the anterior costae meet the anterior margin perpendicularly, as 
is the case in the holotype. In a number of cases the costae which 
approach the ventral margin bend forward at their lower extremities, 
while the anterior costae are uniformly curved right up to the anterior 
margin which they meet at an acute angle; in others the forward 
bend is not evident, though it is to be presumed that with further 
growth it would have made its appearance. The stage at which this 
peculiar feature of the costae appears must vary considerably. In the 
holotype the forward curvature affects the anterior costae as well, so 
that these approach the anterior margin at about a right angle. 

The variations observed appear to be of such a character as would 


Lower Cretaceous Lamellibranchia from Northern Zululand. 337 


be expected within the limits of a species. They apparently have no 
zonal significance, as far as can be judged from the present collection. 

Comparison between T. obesa and T. conocardiiformis (Krauss) 
from the Uitenhage Neocomian.—These two species are very strik- 
ingly similar and certainly closely allied (cf. Plates XX XVIII, XX XIX, 
and XL). They have the same general form and are of about the 
same size, and agree almost exactly in those characters which might 
be taken to be of generic or sub-generic value. The general characters 
of the dorsal area and escutcheon are identical, both having weak 
marginal carinae, which even near the umbones are hardly more than 
folds; both are characterised by the restriction of the pseudo- 
concentric ribbing to the early neanic stage and the complete absence 
of dorsal costellae on the adult area and escutcheon. The ornamen- 
tation of the flank is of the same type and in the juvenile stage all 
but identical. 

The species, however, can be distinguished from each other in 
several respects, the characters being more or less constant for the 
two areas. T. conocardiwformis (see Plate XL) is invariably relatively 
less high and more elongate than T. obesa, with the umbonal region 
less prominent, the antero-dorsal and postero-dorsal margins 
diverging at a larger angle, and the posterior more produced. The 
anterior costae in 7. conocardiuformis are less steeply inclined, and 
are usually more convex towards the posterior, the greater curvature 
and lesser obliquity enabling them to bend round gradually so as to 
meet the anterior margin nearly perpendicularly; the 7. obesa, on 
the other hand, the most anterior costae tend to be steeper and 
straighter, and except in those cases where their lower extremities 
bend forward, meet the anterior margin more acutely. In T. cono- 
carduformis the costae are rounded, crossed by growth lines but 
otherwise smooth, and only sometimes constricted at irregular or 
infrequent intervals; in some individuals the costae are of roughly 
uniform strength for a considerable distance from the umbo; in 
others there are very broad and very shallow concentric furrows 
which correspond to constrictions on the costae, so that the anterior 
costae appear to be made up of a succession of elongate, and less 
commonly oval or rounded, only slightly raised nodes. In I. obesa 
the constrictions tend to be closer together, and the nodes more 
rounded and sometimes more raised, very often giving the costae a 
coarsely beaded character; in 7. conocardiiformis the earlier oblique 
costae within 30 mm. from the umbo may show raised nodes towards 
the anterior margin. The division of the costae into an anterior 


338 Annals of the South African Museum. 


and a posterior series as described by Kitchin (1908, pp. 121, 124), 
a frequent though not an essential character of the Uitenhage species, 
and one which does not appear to mark Krauss’s type, has not been 
noticed in 7’. obesa. 

Comparison with East African Species—The two shells from the 
Trigonia schwarzi-Beds in Tanganyika ascribed by Lange (1914, 
p. 235, pl. xix, figs. 1 a,b) to T. conocarduformis (Krauss) represent 
an unnamed new species which resembles 7’. obesa in certain respects. 
The species is undoubtedly a Megatrigonia from its general form, 
size, costation, and smooth area and escutcheon, the adult shell 
having been well figured by Lange, though somewhat briefly described. 
It appears to be distinguished from TZ. conocardiformis in being 
relatively shorter, and has much the same proportions as T. obesa; 
the anterior costae are finer and more widely spaced than in T. cono- 
carduformis, and less steeply inclined than in 7. obesa. The Hast 
African form must be very closely allied to the two South African 
species and is almost certainly distinct, but as only one specimen 
has been figured it is difficult to make an adequate comparison. 

T. staffi Lange (1914, p. 236, pl. xx, fig. 1), founded on a single 
right valve from the same horizon, is said by its author to be allied 
to T. conocardiformis. The dorsal area and escutcheon were not 
described or figured, and the species may equally well be an elongate 
member of the Scabrae. Lange states that these two species, to- 
gether with 7. rogersi Kitchin (1908, p. 99), belong to the same group, 
but the wide costellate escutcheon of the latter places it among the 
Scabrae. 


Sub-genus Jotrigonia van Hoepen 1929. 


The sub-genus was proposed by van Hoepen (1929, p. 6) for certain 
species from India and South Africa, characterised by the possession 
of smooth V-shaped costae, namely 7. vaw Sharpe and T. stowi 
Kitchin from the Uitenhage Series, and 7. v-scripta Kitchin from the 
Oomia Beds, together with two species from Zululand, Lotrigonia 
crassitesta van Hoepen and I. inconstans van Hoepen. I. crassitesta 
was cited as genotype, and the group was ranked as a genus and 
associated with Megatrigona van Hoepen in the sub-family 
Megatrigoniinae. 

The sub-genus, a brief diagnosis of which has been given also by 
Crickmay (1932, p. 460), may be described as follows:— 

Shell from moderately small to large, considerably longer than 
high, highest towards the anterior end or in front of the middle, the 


Lower Cretaceous Lamellibranchia from Northern Zululand. 339 


posterior much produced and relatively narrow, the anterior rounded 
or produced, moderately convex. Postero-dorsal margin long, 
concave to almost straight; ventral margin very long, convex. 
Flank ornamented with numerous more or less coarse, smooth, 
V-shaped oblique costae, which tend to become irregular or obsolete 
towards the ventral margin or the anterior end. Area moderately 
wide in the adult stage, smooth, separated from the flank by a 
marginal carinal fold which becomes very broad and indistinct 
towards the posterior. Escutcheon long, lanceolate, moderately wide, 
excavated, smooth, separated from the area by a change in slope. 
Neanic stage with smooth concentric ribs, which pass over on to the 
area as transverse ribs; transverse ribs if present at all soon disappear 
from the escutcheon, but persist for some distance on the area; the 
dorsal carinae near the umbones may be sharply marked, linear, or 
represented only by blunt folds; thereis no median carina. Dentition 
strong, the left central tooth markedly bifid. 

Kitchin (1903, pp. 65-67, 115-117; 1908, pp. 110-119) has pointed 
out at some length the grounds for believing that the striking simi- 
larity in adult features between the Indian and Uitenhage species 
is due to homoeomorphy and has distinguished between a “Group 
of T. vau” (including T. stowr) and a “Group of T. v-scripta.”” Con- 
vergence in evolution is indicated in the characters of the neanic 
stage; the former group has a simple youthful ornamentation of 
crowded raised lines which extend from anterior margin to postero- 
dorsal margin without a break, while area and escutcheon are hardly 
defined, the marginal carina being a blunt fold and the inner carina 
being more or less obsolete; in the Group of T. v-scripta, on the other 
hand, the youthful sculpture is more differentiated, the ribs on the 
flank are fewer and more widely spaced and bend obliquely forward 
at the well-defined marginal carina, to terminate at the delicate 
linear ridge which represents the inner carina. Nevertheless it 
should be pointed out that the distinction made by Kitchin, though 
of importance, does not necessarily indicate descent from very 
remotely related ancestral stocks, since the youthful ornamentation 
in the two groups is in a broad way of the same type. The sub- 
generic term Jotrigonia can therefore be used for both groups. The 
genotype, I. crassitesta van Hoepen (1929, p. 7, pl. u1, figs. 4, 5; pl. in, 
figs. 1, 2), apparently belongs to the Group of T. vau. 

It is rather curious that van Hoepen, who set out to classify the 
Zululand Trigoniae on phylogenetic lines (ibid.; p. 3), included both 
T. vau and T. v-scripta in Iotrigonia without any mention of the 


340 Annals of the South African Museum. 


remarkable convergence shown by these two species and so well 
described by Kitchin. 

Kitchin (1903, pp. 65-78) has stated that the costation differs 
materially from the analogous sculpture which characterises the 
members of the Jurassic Undulatae of Europe, and that the general 
resemblance is of the remotest kind. The Undulatae are more or 
less palpably derived from the Clavellatae, doubtless along more 
than one line of descent, but the ancestry of the Indo-African 
Neocomian group is by no means certain. Crickmay (1932, p. 457) 
has proposed a genus Vaugonia for a member of the Undulatae, and 
Dietrich (1933, p. 33) has briefly discussed the convergence shown 
by these V-costate species. There appears to be some justification 
for associating Lotrigonia with Megatrigonia, since both are charac- 
terised by a primitive concentric ornament in the neanic stage, by 
smooth area and escutcheon in the adult stage, and obsolescent 
marginal and inner carinae, but the adult costation is of quite a 
different character in the two sub-genera; a remote common ancestor 
is possible, as originally suggested by Kitchin (1908, p. 122). 

The two species described by van Hoepen (1929, pp. 7-9) from the 
Umsinene River in Zululand are strikingly similar to the two species 
from the Uitenhage Series. T. crassitesta (van Hoepen) appears to 
be close to T. stowi Kitchin (1908, p. 115, pl. vi, figs. 4, 5; pl. vu, 
fig. 1) and JT. inconstans (van Hoepen) is very similar to but con- 
siderably larger than 7. vaw Sharpe (Kitchin, 1908, p. 110, pl. vi, 
figs. 1-3). Both species were collected at the base of the succession, 
from van Hoepen’s Ndabana Beds, where they were associated with 
Megatrigoma obesa, and a member of the group of the Scabrae 
reminiscent of the Uitenhage T. kraussi though distinct from it. 
The Ndabana Beds were stated to be Albian by van Hoepen (1926, 
p. 222) in a preliminary account, but the Trigoniae which have been 
collected belong to groups which are more characteristic of the 
Neocomian. 


Trigoma (Lotrigonia) haughtoni sp. nov. 
(Plate XLI, figs. 1-4.) 


Material._—The species was collected by Dr. 8. H. Haughton and 
Dr. A. W. Rogers from two neighbouring localities, in both cases 
in association with T. (Megatrigonia) obesa and T. (Steonmanella) 
hennigi. There are five pieces of shelly limestone from MI on the 
Myesa tributary of the Pongola River, including portions of 10 
detached valves, both right and left valves being represented, as 


Lower Cretaceous Lamellibranchia from Northern Zululand. 341 


well as a right valve from M3. From ZA on the Mfongosi stream 
3 specimens. Several of the valves are more or less complete and 
in a good state of preservation. 

Holotype.—The holotype is the largest of four specimens associated 
on the same slab from MI, and is a fairly complete and well-preserved 
right valve. Two of the associated specimens, onea right valve and the 
other a left valve, show the neanic stage and the dorsal area very finely, 
and may be cited as paratypes. I have not seen the interior of the 
valves. The postero-dorsal part of the holotype appears to be damaged. 

Description of the Species.—Shell moderately small, considerably 
longer than high, with the umbo situated at about a third of the 
length of the shell from the anterior end, umbonal region rather 
prominent, anterior relatively short and rounded, posterior produced 
and narrower. Shell in the adult stage moderately and more or less 
evenly convex; the younger shell relatively more convex. Anterior 
margin broadly convex, the greatest curvature at or about the 
middle, merging quite gradually into the long convex ventral margin; 
postero-dorsal margin moderately long, gently to strongly concave 
for some distance behind the umbo, meeting the ventral margin in a 
well-rounded posterior termination. Umbo well incurved. 

Area and escutcheon taken together wide, more or less crescentic, 
broadening rather rapidly at first and very steeply inclined, when 
traced posteriorly gradually becoming a little narrower and less 
steeply inclined. Marginal carina represented by a raised, very 
convex, rounded, smooth fold, which gradually becomes wider and 
relatively less raised when traced posteriorly; the marginal carinal 
fold pursues a course which is markedly concave towards the postero- 
dorsal margin, and also concave upwards when the valve is seen in 
lateral view; from about 5 mm. to about 20 mm. from the umbo the 
lower side of the fold is more or less distinct by reason of its con- 
spicuously raised character, and the costae of the flank commence 
immediately below it; traced posteriorly the fold broadens con- 
siderably and is less distinctly limited from the flank. Area narrow, 
narrower than the escutcheon except near the posterior extremity; 
up to 5 mm. from the umbo the area is very steeply inclined and 
slightly concave; beyond 5 mm. from the umbo the area gradually 
becomes deeply and evenly concave in cross-section, merging below 
into the convex marginal fold and on the upper side meeting the 
escutcheon in a raised crest; the median longitudinal furrow is not 
distinguishable in the general concavity of the area; at the posterior 
end the marginal fold has broadened out so as to make up the greater 


342 Annals of the South African Museum. 


part of the width of the area, this broad, gently convex part being 
succeeded upwards by a much narrower concave part. The concave 
area meets the wider but equally concave escutcheon in a broadly 
V-shaped, sharp-crested inner carina, which pursues a course at 
first markedly concave towards the postero-dorsal margin. Both 
the marginal carinal fold and the inner carina gradually become 
less concave towards the postero-dorsal margin as they are traced 
backwards; both appear concave upwards when the shell is seen in 
side view. Escutcheon broad, evenly and deeply concave from the 
raised inner carina to the raised upper margin, crescentic in shape. 

In the neanic stage, up to 5 mm. from the umbo, the ornamentation 
consists of numerous concentric ribs which extend from the anterior 
margin to the marginal carinal fold in a uniform curve, meeting the 
fold almost perpendicularly. At from 2 mm. to 5 mm. from the 
umbo the ribs on the flank are raised, rounded, and separated by 
wider concave interspaces. Up to about 2 mm. from the umbo the 
ribs on the flank curve slightly downwards on crossing the carinal 
fold and then bend upwards so as to cross the area very obliquely. 
Between 2 mm. and 5 mm. from the umbo the ribs on the flank are 
not continuous with the corresponding ribs on the area, the carinal 
fold being smooth except for growth lines; the ribs on the area are 
very oblique, slightly less raised, and closer together than the corre- 
sponding ribs on the flank. Beyond 5 mm. from the umbo the 
coarse oblique costae of the posterior series make their appearance, 
but the primitive oblique ribbing of the area persists to about 10-12 
mm., the ribs being less close together than at an earlier stage and 
becoming confined to the centre of the concave area. Beyond about 
5-7 mm. from the umbo the ribs on the area fail to reach the inner 
carina. At a very early stage the ribs probably pass over on to the 
escutcheon (?), but at 4-8 mm. from the umbo there are only traces 
of very fine oblique riblets just above the inner carina, but these do 
not persist when traced towards the upper margin. The greater 
part of the area and escutcheon is smooth. The ribs appear to have 
been smooth, but the inner carina bears traces of very imperfectly 
developed nodes up to about 15 mm. from the umbo. 

The adult shell is ornamented with two series of oblique costae so 
arranged as to give a V-costate appearance, but the anterior and 
posterior series do not coincide in numbers of costae. The adult 
type of ornamentation appears abruptly at about 5 mm. from the 
umbo when the posterior series makes its appearance; a concentric 
tib which shows no sign of being divided into a posterior and an 


Lower Cretaceous Lamellibranchia from Northern Zululand. 343 


anterior limb is succeeded immediately by the two adult series of 
inclined costae. The primitive concentric ribbing, however, grades 
into the anterior series of costae, which are directed steeply postero- 
ventrally and cross the growth lines at an angle of 15-25°. The 
anterior costae are rounded, smooth, moderately elevated, and are 
about twice as numerous as the coarser posterior series; they pursue 
a nearly straight course from the anterior margin, but are very slightly 
concave to the postero-dorsal margin. The line of junction of the 
anterior and posterior series is nearly vertical, but very steeply 
inclined backwards. The posterior costae are coarser and radiate 
downwards from the margin of the area, the 4 or 5 costae nearest 
the anterior being directed very steeply forwards so as to abut against 
the anterior series; these are succeeded posteriorly by vertical costae, 
and the shorter costae which follow are directed very steeply back- 
wards. The posterior costae are rounded, convex, smooth, separated 
by concave interspaces of about the same width as themselves; the 
4 or 5 nearest the anterior are very wide and coarse from their junction 
with the anterior series until close to the area, but become narrow 
close to the marginal carinal fold; traced posteriorly the costae 
become progressively narrower and shorter. The costae fail to reach 
the ventral margin in the adult shell, persisting to about 25 mm. 
from the umbo in the anterior part. 
All parts of the shell are crossed by growth lines. 
Dimensions, in millimetres: 


Holotype. Paratype 

(left valve). 
Length . : 45 +24 
Meee se el 19 
Thickness , : a. 7 + 5 


Comparison with other South African Species.—This fine little 
species 1s quite distinct from each of the four South African species 
of Lotrigonia described up to the present, and does not appear to be 
very closely allied to any of them. The considerably smaller size 
and rounded anterior serve to distinguish 7. haughton: at a glance 
from the more elongate, anteriorly produced T. stow: Kitchin (1908, 
p. 115, pl. vi, figs. 4, 5; pl. vii, fig. 1) and Lotrigonia crassitesta van 
Hoepen (1929, p. 7, pl. ii, figs. 4, 5; pl. ii, figs. 1, 2). In outline 
there is some resemblance to the considerably larger I. anconstans 
van Hoepen (1929, p. 8, pl. iii, figs. 3, 4; pl. iv, figs. 1, 2) and to 
T. vau Sharpe (Kitchin, 1908, p. 110, pl. vi, figs. 1-3). The resem- 
blance, based on size and outline only, is greatest to 7. vau, 


344 Annals of the South African Museum. 


but the species can be distinguished in a number of important 
respects. 

T. vau is larger than T. haughton. In the former both the anterior 
and posterior series of costae are inclined much less steeply, and 
consequently meet in a considerably larger angle; the line of junction 
of the two series 1s directed obliquely downwards, and is not as 
nearly vertical as in T. haughtom. In T. haughtoni there are about 
twice as many costae in the anterior series as in the posterior series, 
where they meet in the line of junction, whereas in T. vau the numbers 
are nearly the same, the majority of the posterior costae passing 
with a sharp V into the anterior costae, there being only one or two 
additional costae on the anterior side. In T. vau the posterior costae 
are all directed forwards, while in 7. haughtoni the most posterior 
of the costae are directed steeply backwards. In T. haughtoni the 
marginal carinal fold is more convex and raised than is the case in 
T. vau, the area is relatively wider, the escutcheon is relatively 
narrower and smaller, the inner carina more strongly developed, 
and the ribbing of the area more persistent. 

As pointed out above, Kitchin has distinguished between a “Group 
of T. vaw” and a “Group of T. v-scripta,” basing the distinction on 
the characters of the neanic stage. The former group includes the 
two Uitenhage species T. vau and T. stowr, both characterised by 
simple concentric ribbing in the neanic stage and obsolescence of the 
inner carina; in these species the ribs are linear and crowded, and 
pass on to the area without interruption. The character of the 
youthful stage in T. haughtoni is such as to suggest relationship with 
the group of 7. v-scripta rather than with the two Uitenhage species 
T. vau and T. stowi, noteworthy being the fact that in T. haughtona 
the concentric ribs bend rather sharply upwards at a well-defined 
carinal angle as they do in the Indian species, whereas in 7. vau and 
T. stowi they pass over on to the area with only a slight change in 
direction, the carinal angle being much rounded. The fact that the 
inner carina is raised and sharp-crested supports reference to the 
Indian group, since in the Uitenhage species the inner carina is a 
weak fold. On the other hand, the concentric ribs are as numerous 
as in T. vau, the Indian species being characterised by the possession 
of a few relatively coarse ribs within 5 mm. from the umbo. 

Comparison with other Species.—There is a certain resemblance in 
general features to T. v-scripta Kitchin (1903, p. 70, pl. vu, figs. 
6-8; pl. vin, figs. 1-3) from the Oomia Beds of Cutch, but the species 
differ from one another in details of form and ornament. A note- 


Lower Cretaceous Lamellibranchia from Northern Zululand. 345 


worthy distinction is the considerably larger number of concentric 
ribs in the neanic stage in the new species. T. haughtoni has greater 
juvenile convexity, and other points of distinction are the slightly 
greater obliquity of the line of junction of the anterior and posterior 
series of costae, the wider escutcheon and more curved inner carina, 
the sharp demarcation of escutcheon and area, the steep backward 
slope of the most posterior costae, and the rather shorter anterior. 
The species are undoubtedly distinct, but an actual comparison of 
specimens would be necessary for an exhaustive and adequate 
comparison. 

T. dubia Kitchin (1903, p. 67, pl. vu, figs. 3-5) from the same 
formation as the above appears to be relatively longer and broader (?) 
in the posterior region, and the anterior series of costae is concentric 
or almost so. Dietrich (1933, p. 33, pl. 11, figs. 45, 46) has identified 
with the Indian species some very imperfect shells, from the 7. smeei 
beds in Tanganyika, which resemble the new species in the character 
of the costation, and in which the anterior series of costae is oblique 
and probably not parallel to the ventral border; the change from 
primitive to adult ornamentation is, however, not as sudden as in 
the new species, there being several concentric ribs more or less 
sharply bent in the middle. T. dubia is imperfectly known, and it 
is very doubtful whether the East African shells represent the same 
species. 

I have not been able to see the paper containing the description 
and figures of the Hast African species 7. kuhni G. Miiller, said by 
Kitchin (1908, p. 113) to be of Neocomian age. Judging from 
Kitchin’s remarks, the species appears to have been founded on 
imperfect material. 

T. heterosculpta Stanton (1901, p. 20, pl. iv, figs. 16-18) from the 
Lower Cretaceous of Patagonia is characterised by the fact that the 
anterior costae terminate against a single vertical posterior rib, and 
differs from the new species in several other respects. 


Sub-genus Stexnmanella Crickmay 1930. 
(=Transitrigoma Dietrich 1933.) 


The species described below is a typical example of the very distinct 
Neocomian group of Trigoniae long known as the Pseudo-quadratae, 
a name bestowed by Steinmann (1882) to include two southern 
species, 7’. transitoria Steinmann from South America and 7. herzogi 
(Goldfuss) from the Uitenhage Series in South Africa. Several species 


346 Annals of the South African Museum. 


have since been added from South America, as well as T. holubi 
Kitchin from the Uitenhage Series, 7. mamillata Kitchin from the 
Oomia Beds of Cutch, and T. hennigi Lange from Hast Africa. In 
America the group has been recognised as far north as Texas, where 
the species T. vyschetzkii Cragin occurs with certain other Lamelli- 
branchs of Lower Cretaceous affinities in the so-called Malone Jurassic 
Formation; Kitchin (1926) has set forth very fully reasons for believ- 
ing that the Lamellibranchs concerned came from Neocomian beds 
included in that formation. The Pseudo-quadratae have been dis- 
cussed by Kitchin in several papers (1903, p. 98; 1908, p. 108), and 
were thought by that author to be confined to the Neocomian (1926, 
1929). 

To this well-marked group two generic names have recently been 
applied, Steonmanella Crickmay (Nat. Mus. Canada, Bull. 63, 1930), 
and Transitrigonia Dietrich (1933, p. 36). The designated types are 
very closely allied species, and T'ransitrigonia (type T. transitoria 
Steinmann) is therefore a synonym of Stevnmanella (type T. holubi 
Kitchin). A brief diagnosis of Steonmanella has been given by 
Crickmay (1932, p. 458) in his “Contributions towards a Monograph 
of the Trigoniidae.” 

The sub-genus may be described as follows :— 

Shell large, massive, quadrate in outline, compressed. Umbones 
nearly terminal, weakly incurved. Anterior more or less vertically 
truncate, the anterior margin sloping steeply downwards from the 
umbones and meeting the long straight postero-dorsal margin nearly 
perpendicularly. Posterior high, the posterior margin relatively long. 
Area very broad, forming with the flank a gentle uniform convexity, 
with a weak longitudinal furrow above the middle; ornamented 
towards the umbones with three longitudinal rows of tubercles, re- 
presenting the marginal and inner carinae and a median carina just 
below the furrow; ornamented beyond the neanic stage with growth 
lines which may at some stage pass into coarse, crowded, smooth, 
concentric costellae. Escutcheon narrow, very elongated, ornamented 
with irregular oblique rows of coarse tubercles. Flank beyond the 
neanic stage with widely spaced, oblique, coarsely tuberculate costae, 
or rows of coarse tubercles. Harly neanic stage with relatively strong 
concentric nodular ribs, which pass from the anterior margin across 
the flank on to the area and escutcheon. Dentition massive, the left 
central tooth narrow. 

The characters shown by the neanic stage indicate descent from 
the Jurassic Clavellatae (Kitchin, 1908, p. 109). 


Lower Cretaceous Lamellibranchia from Northern Zululand. 347 


Trigoma (Steonmanella) hennigr Lange. 
(Plate XLIII, figs. 1-3; Plate XLIV, fig. 1.) 


1914. Trigonia Hennigr Hanes: Tendaguruschichten, p. 238, taf. xix, 
fig. 3. 

1914. Trigonia transitoria Steinmann: Lange, ibid., p. 237, taf. xix, 
fig. 2. 

1933. T. (Lransitrigonia) hennigi Lange: Dietrich, p. 37. 

Materval—Twelve specimens identified with this East African 
species were available for study, of which eleven were collected by 
Dr. 8S. H. Haughton and Dr. A. W. Rogers along tributaries of the 
Pongola River, and one by H. F. Frommurze at an unrecorded horizon 
on the Mfongosi stream. The later collection of eleven specimens 
was found at the following localities: Myesa stream, MI, 2 juvenile 
specimens; Mfongosi stream, ZA, 6 specimens; Lombagwenya 
stream, one specimen from each of the localities L9, L12, L14. The 
specimen (S.A.M. 10786) collected by H. F. Frommurze is very 
finely preserved, and it is probable that it is from ZA. The majority 
of the shells have the valves ciosed, and though complete are not well 
preserved. The interior has only been seen in part. Dr. 8S. H. 
Haughton also recorded the species at locality L4. 

Description of the Zululand Shells. (1) Haternal Features.—Shell 
large, massive, oblong in outline, longer than high, not much inflated. 
Dorsal margin long, more than two-thirds of the length of the shell, 
nearly straight. Anterior margin very gently convex, forming about 
a right angle with the dorsal margin, at first vertical or very nearly 
so, then curving very gradually backwards, merging into the convex 
ventral margin about a much-rounded antero-ventral angle. Dorsal 
margin meeting the posterior margin in a very obtuse angle. Posterior 
margin very broadly convex, very obscurely angulated above the 
middle and meeting the ventral margin in a much-rounded obtuse 
angle. 

Umbones small, pointed, slightly incurved and almost terminal 
at the antero-dorsal angle. Anterior more or less truncated, with a 
narrow flattened or only slightly convex anterior face almost at right 
angles to the plane of the valves, well demarcated from the flank 
not only by the difference in slope but by the absence of costae. 
Inflation of valves relatively weak, the greatest Se cea above the 
middle of the valves and towards the anterior. 

Dorsal area very wide, marked off from the flank by the absence 


348 Annals of the South African Museum. 


of the coarse oblique costae. Marginal carina represented only by 
a very broad gentle curvature of the valve surface. The area is 
traversed longitudinally by a well-marked shallow furrow, which 
persists from the umbonal region to the posterior margin, dividing 
the area into a narrow upper and a broader lower part, the lower 
part being 2-3 times as broad as the upper. The escutcheon is 
rather narrow and elongate, and is separated from the dorsal side 
of the area by an inner carina, which is well marked within 20 mm. 
of the umbo; the inner carina when traced posteriorly is gradually 
replaced by an inconspicuous fold that persists to the postero-dorsal 
angle. The escutcheon is not much excavated near the umbones, 
and is slightly convex in the posterior half. There is a short, deep, 
very narrowly cordate lunule well demarcated from the anterior face 
by an abrupt change of slope. 

(2) Internal Features.—Interior of the valves not seen, except for 
part of the sub-umbonal region of a large left valve, in which there 
is a very massive, relatively narrow central tooth, projecting to about 
12 mm. above the margin, oblique, coarsely grooved on the almost 
vertical sides, and deeply channelled above; there is also a relatively 
strong anterior tooth, narrow, slightly oblique, and close to the 
anterior margin. 

(3) Ornamentation.—Flank of the shell ornamented with about 
8-10 very coarsely tuberculated costae, which pass gradually into 
finer concentric costae within about 8 mm. from the umbones. Costae 
oblique, from nearly straight to distinctly or even markedly concave 
towards the anterior. In some specimens the costae behind the 
middle of the flank bend forwards rather abruptly at some distance 
from the ventral margin. In the umbonal region the costae are 
short and directed antero-ventrally, in the mid-region they are more 
nearly vertical, and they become progressively shorter posteriorly 
as the dorsal area widens, the last one or two being directed slightly 
backwards. ; 

The costae are characterised by bearing very large, prominent, 
rounded tubercles at frequent and fairly regular intervals; the costae 
themselves are not much elevated, but are given a strong appearance 
by the very prominent swollen tubercles. The tuberculation may 
become a bit irregular towards the ventral, particularly the antero- 
ventral, border. Anteriorly the costae cease abruptly at the angle 
limiting the anterior face, so that in anterior view the narrow anterior 
face appears to be bounded by a row of coarse tubercles. The costae 
are widely separated by concave interspaces, which are much wider 


Lower Cretaceous Lamellibranchia from Northern Zululand. 349 


than the costae in the ventral half. The costae end somewhat 
abruptly at the margin of the dorsal area, near or a little below 
the obscure carinal convexity. 

The area in the adult stage is concentrically striate and devoid 
of tubercles, but there are indications of two longitudinal rows of 
low coarse tubercles which persist to about 30 mm. from the umbo, 
one row representing the obscure marginai carina and one row re- 
presenting an obscure median carina just below the longitudinal 
furrow. ‘The inner carina carries a row of rather prominent rounded 
tubercles for some distance, but these disappear before the posterior 
margin is reached. ‘The escutcheon bears several indistinct and very 
oblique rows of moderately large elongate tubercles. 

All parts of the surface are traversed by relatively fine growth lines 
of varying strength. There is little indication in most specimens 
that the concentrically striate area changes in character with age; 
the coarse concentric costellae which characterise the greater part, 
or at least the later part of the area in other species, do not make 
their appearance until a very late stage. In 8.A.M. 10786 concentric 
costellae are certainly not developed at 100 mm. from the umbo; 
the posterior border, however, is too damaged for one to be quite 
certain that they do not appear at all. In the largest specimen, a 
weathered right valve from horizon L14, there are indications of con- 
centric costellae at about 120 mm. from the umbo, but I have not 
been able to trace these across the area owing to the poor state of 
preservation of the specimen. 

Within about 8 mm. from the umbo the costae are relatively pro- 
minent, nodular, and concentric, and are continued across the slightly 
raised marginal carina over the area at least as far as the raised inner 
carina. On crossing the marginal carina these costae form a distinct 
V, the apex directed ventrally. Beyond about 5 mm. from the umbo, 
the concentric costae on the area break up into three nearly contiguous 
tubercles, which at about 20 mm. are distinguishable as the three 
divergent longitudinal rows of tubercles representing marginal, 
median, and inner carinae. 

(4) Dimensions.—Of the following specimens, whose dimensions 
are recorded in millimetres, 1 is S.A.M. 10786; 2-6 are from 
ZA; 7 and 8 are from MI; 9, 10, and 11 are from L9, L12, and 
L14 respectively. 

am, @. ©. G. GoGe wo Gy Oe (Ona: 
Length . : . 116 108 104 112 101 119 92 60 125 116 126 


Height . : Gr aii owed: tl Sl 728 45° 92) Sl SF 
Thicknessofonevalve 22 21 Opel 6ee Flite 22) Le 2 S232 


VOL. XXXI, PART 3. 29 


350 Annals of the South African Museum. 


Comparison with Uitenhage. Species.—The Pseudo-quadratae have 
hitherto been represented in the South African Cretaceous by two 
species from the Sundays River Beds, the age of which has been 
shown to be Upper Valanginian by L. F. Spath (1930, p. 132). These 
are T. herzogi (Goldfuss) and the less common 7. holubi Kitchin, the 
latter the genotype of Steinmanella; both species have been described 
and figured by Kitchin (1908, pp. 101-110). 

The shells in the present collection are quite distinct from the two 
Uitenhage species, though clearly allied to both. I have been able 
to compare them directly with the examples figured by Kitchin, 
including the holotype of 7. holubi, as well as with several other 
specimens from the Uitenhage division. The Zululand shells are 
of roughly the same size as the two Uitenhage species, and agree 
in having robust, little inflated, sub-quadrate valves, with wide area 
and coarsely costate flank, besides being strikingly similar in the 
general character of form and sculpture. They differ, however, from 
both species in outline, in many details of ornamentation, and notably 
in the nature of the area in its adult stage. In so far as outline and 
costation are concerned they represent a species intermediate in 
character between the two Uitenhage species, exhibiting a combina- 
tion of features which invites comparison with both of them. 

The Zululand shells have the same general proportions as T. holubi, 
and are distinctly shorter than T. herzogi, which is the most elongate 
species in the sub-genus. The ratio of length to height is 1: 0-7 in 
T. holubt and in the Zululand shells, and 1: 0-55 in T. herzogi. In 
this respect they approach the relatively short 7. transitoria and 
other species. 

In the character of the anterior part of the shell the Zululand 
specimens are distinct, but features of both Uitenhage species are 
recalled. In J. herzogi the anterior margin is broadly and evenly 
convex from the umbones downwards, and merges quite gradually 
into the ventral margin, with little or no sign of anterior truncation 
and without a flattened frontal face; the general trend of the anterior 
margin, which at first is gently convex forwards, is steeply downwards 
and backwards from the umbones. In 7. holubi, on the other hand, 
the anterior margin is very slightly convex to almost straight, and 
is directed vertically from the umbones or inclined forwards at a 
steep angle to meet the ventral margin in a well-rounded but distinct 
angle; the anterior is markedly truncate, and a narrow, flattened 
frontal face is developed. In the holotype of 7. holubi, 2.e. the only 
specimen figured by Kitchin (1908, p. 103, pl. iv, figs. 2, 2 a), the 


Lower Cretaceous Lamellibranchia from Northern Zululand. 351 


anterior margin is inclined forwards at a steep angle, but in other 
specimens it is directed almost vertically. In the Zululand shells 
the anterior margin is gently and evenly convex, with about the same 
curvature as in Tf. herzogi, and the general trend of the anterior 
margin is very steeply downwards and backwards; the anterior 
margin meets the ventral margin in a very much rounded, obscure, 
or almost distinct angle; in the above features these shells resemble 
T. herzogi very closely, though in the latter there is no sign of the 
antero-ventral angle. On the other hand, the anterior is markedly 
truncate in these shells, a narrow, flattened, or only slightly convex 
frontal face being developed as in T. holubi; but the frontal face is 
ornamented only with very oblique growth lines, whereas in 7. holubi 
there are coarse folds which blend with the flank costae. 

In the character of the costae on the flank of the valves the Zulu- 
land specimens are again distinct, features of both Uitenhage species 
being recalled. In TZ. herzogi the flank is ornamented with almost 
straight, regular, oblique, coarsely tuberculate costae, with the 
tubercles close to one another; the costae become somewhat irregular 
close to the ventral border only; the costae close to the anterior 
margin are inclined steeply forwards, in the mid-region they are 
directed vertically, and in the posterior half are inclined steeply 
backwards. In T. holubi, on the other hand, the costae are much 
less regular, and in the mid-region are broken up into widely separated 
coarse tubercles; the rows of tubercles or costae are regularly curved 
forwards when traced downwards from the margin of the area, and 
are never directed backwards; towards the ventral border the 
tubercles are crowded, and a tendency to elongation parallel to the 
shell margin exhibits itself, so that with senility crowded growth 
ridges are dominant at the ventral border. In the Zululand shells 
the costae have the general character of those of T. herzogi, 1.e. they 
are regularly tuberculate with close-set and evenly spaced tubercles; 
but the costae are either nearly straight or curve forwards when 
traced downwards from the area, asin T. holubi. It should be pointed 
out, however, that while the forward inclination of the costae is 
characteristic of 7. holubi the costae are not necessarily as irregular 
or the tubercles in the mid-region as widely spaced as in the holotype 
of that species. I have before me a specimen (A.M. 2356) from 
the Sundays River Beds which, while agreeing with the type of 
T. holubi in other respects, has fairly regular costae in the mid- 
region with the tubercles moderately close to one another; as 
the specimen differs from the holotype also in that the anterior 


302 Annals of the South African Museum. 


margin is vertical, it is, however, possible that it belongs to a 
distinct variety. 

It is in the character of the area, however, that the Zululand shells 
are strikingly different from both Uitenhage species. In the Pseudo- 
quadratae the early neanic stage of the area is crossed by transverse 
ribs continuous with those of the flank; these soon pass into a 
“trituberculate” stage in which the area is characterised by bear- 
ing three longitudinal rows of tubercles representing marginal, 
median, and inner carinae, the area being otherwise smooth 
except for growth lines. At some distance from the umbo coarse 
crowded concentric costellae make their appearance and these 
characterise the adult stage. As Kitchin (1908, pp. 108-110) has 
pointed out, the Pseudo-quadratae pass through a similar sequence 
of developmental phases, but these phases are not reached by all the 
species at the same time. In T. herzogi the “trituberculate” stage 
may be seen to pass into the “‘costellate” stage by the progressive 
transverse elongation of the tubercles in the three sets, and the change 
takes place before mid-growth. In T. holubi the change takes place 
after mid-growth and is more abrupt, but the three rows of tubercles 
persist until the “costellate” stage is reached. In the Zululand 
shells the marginal and median rows of tubercles die out at some 
time before mid-growth, while the inner row is more persistent, but 
the “trituberculate” stage is not replaced by a “‘costellate” stage, 
the greater part of the area being merely transversely striate with 
growth lines; the growth lines are hardly stronger on the area than 
they are in the intercostal interspaces on the flank. In 8.A.M. 
10786 the marginal and median rows of tubercles persist to about 
40-50 mm. from the umbones and the inner row to about 55 mm., 
while costellae have not made their appearance even at 100 mm., 
about 10 mm. from the damaged posterior border. In the larger 
weathered specimen from Ll4 there are, however, indications of 
costellae on the lower side of the area at about 120 mm. from the 
umbo, and though their character is a little uncertain it would seem 
that they do make their appearance with senility. This delay in the 
appearance of the costellae is one of the most noteworthy features of 
the Zululand shells, and they would appear therefore to be more 
primitive—though not necessarily earlier in age—than the two Uiten- 
hage species and their allies in 8. America and India, in all of which 
racial degeneracy is in evidence (Kitchin, 1908, p. 110). 

Comparison with Hast African Species and with T. transitoria 
Steinmann.—The Pseudo-quadratae have been recorded from the 


a 


Lower Cretaceous Lamellibranchia from Northern Zululand. 353 


Trigoma schwarzt Beds in Tanganyika by Lange (1914, pp. 237-239), 
who described a new species, 7’. hennigi, as well as certain shells which 
he identified with the very widespread South American species, 
T. transitoria. One is inclined to be suspicious of the identification 
of an Kast African Trigonia with a South American species, and the 
suspicion is perhaps strengthened by the absence of that species from 
the Uitenhage Series. A comparison of the set of shells before me 
with the descriptions and figures given by Lange has led me to 
conclude that Lange’s 7. transitoria should be included in T. hennigi, 
and to accept identification of the Zululand shells with that species. 

Lange had four specimens before him, from three separate localities, 
and three of these specimens were identified with 7. transitoria. The 
fourth specimen, which is from a different locality to the others, is a 
right valve, and was made the holotype of T. hennigi Lange in spite 
of the fact that the upper part of the flank is considerably denuded, 
as is shown clearly enough in the good photograph provided (2bid., 
pl. xix, fig. 3). The type of T. hennigi is said by Lange to differ 
from the other specimens in that the rows of tubercles are straight, 
the ventral margin is less convex, and the inflation of the valves 
extends further back, and it is stated that there is a smooth region 
completely devoid of tubercles below the dorsal area. The principal 
feature which no doubt prompted Lange to recognise a distinction, 
is the fact that the costae or rows of tubercles are traceable upwards 
from the ventral margin for not much more than one-third of the total 
height of the valve, and are not seen to reach the margin of the area 
in the anterior half of the shell. The figures given fail to bring out 
any real difference in the curvature of the ventral margin, and the 
slight difference in the inflation mentioned can hardly in itself be 
of much significance. The photograph of the type, however, clearly 
shows that the whole of the upper two-thirds of the anterior half of 
the valve has suffered severely from either solution of calcite or wear, 
and that the costae cease as this denuded region is approached. 
I have no doubt that prior to weathering the costae passed up to the 
border of the area, as in all other species in the sub-genus, and it is 
highly probable that they were gently curved. The recognition of 
this fact removes any real reason for separating from 7’. hennagi the 
other East African shells studied by Lange (ibid., p. 237, pl. xix, 
fig. 2), for the figures given are in general agreement in other respects, 
the outlines being almost identical, and the ornamentation of the 
same type throughout in so far as it can be discerned. 

The East African species 7. hennigi is then a well-marked type 


304 — Annals of the South African Museum. 


characterised by the forward curvature of the costae and the slightly 
receding anterior margin. The anterior margin is very gently convex 
and almost vertical, but traced downwards becomes considerably 
more convex as it passes back into the ventral margin.’ The costae 
are slightly curved with forward convexity, but their lower portions 
may be almost straight, and the shorter posterior costae are not 
directed backwards. These features as well as the size and pro- 
portions of the valves very strongly suggest identity with the Zululand 
shells, described above. The type of 7. hennigi is further said to 
have a completely smooth frontal face perpendicular to the flank, 
and the area in the adult stage is said to be apparently ornamented 
only with growth lines, points which characterise the Zululand shells 
also, though it should be mentioned that on Lange’s figure there is 
some slight indication of concentric corrugation towards the posterior. 
Lange’s “7. transitoria” is only briefly described, and it is stated 
that the broad area is transversely ribbed; on the single figure given 
the area is unfortunately very worn, but it would appear that coarse 
costellae are not developed, and the transverse ribs mentioned are 
almost certainly no more than growth lines. I can find no grounds 
for separating the Zululand shells from 7. hennigi, though it is to 
be regretted that the type of the latter—as well as the other East 
African shells—are not sufficiently well preserved for a complete 
comparison to be made. 

From T. transitoria Steinmann (1882), which has been described 
from the Neocomian of South America from Patagonia to Bolivia, 
T. hennigi differs in several respects. The former is characterised 
notably by the larger number of costae and the coarse crowded 
transverse costellae which characterises the area for the greater part 
of its length. The difference in character will be readily appreciated 
by comparing the figure of the finely preserved Zululand shell, 
S.A.M. 10786 (pl. xlii, figs. 1-3), with published figures of 7. transi- 
toria (refs. in Lange, 1914, and Kitchin, 1908), and with Lange’s 
‘“T. transitoria”’ (Lange, 1914, pl. xix, fig. 2). 

Comparison with other Species.—The notable absence of costellae 
on the area, except perhaps with senility, serves to distinguish 
T. hennigi from the rather smaller 7. mamullata Kitchin (1903, p. 100, 
pls. ix, x) from the Oomia Beds of Cutch, in which costellae appear 
at an early stage and are remarkably coarse; in other respects the 
species are very similar, though the anterior face is less clearly marked 
off in the Indian species. The absence of costellae serves also to 
distinguish the species from T. vyschetzkw Cragin (1905, p. 56, pls. vi, 


Lower Cretaceous Lamellibranchia from Northern Zululand. 355 


ix) from Texas, and the South American 7. neuquensis Burckhardt 
(see Kitchin, 1908, p. 108), both closely allied to 7. transitoria. 


Sub-genus Rutitrigonia van Hoepen 1929. 


Three generic or sub-generic terms have recently been proposed 
for concentrically ribbed Trigonae from the Indo-African region, 
viz. Rutitrigonia van Hoepen and Pleurotrigonia van Hoepen (1929, 
pp. 31-34) for species from Zululand, and Indotrigonia Dietrich 
(1933, p. 30) for the Indian and Hast African T. smeei J. de C. 
Sowerby. ‘The type species of these three groups have in common 
the possession of a flank ornamentation of coarse smooth concentric 
ribs as in the Jurassic Costatae, but are distinguished from the 
latter in that dorsal ornamentation of a longitudinal character, if 
present at all, is confined to the neanic stage, the area through the 
period of middle growth and maturity being either smooth or 
traversed by concentric costellae. In the type of Indotrigonia, as 
has been pointed out by Kitchin (1903, pp. 39, 42-44, 113) and 
Dietrich (loc. cit.), there is fine evidence of descent from the Costatae 
in the differentiated juvenile ornament of smooth concentric ribs 
ceasing posteriorly at a narrow ante-carinal space, beaded dorsal 
carinae, and fine longitudinal granular ridges on the area. In the 
type of Pleurotrigonia the evidence points likewise to Costate ancestry, 
as is pointed out elsewhere in this paper; the juvenile stage having 
concentric ribs, ante-carinal space, and sharp marginal carina, and 
longitudinal ornament on the dorsal area. Indotrigonia and Pleuro- 
trigoma are, however, readily distinguishable on adult characters; 
the smooth area, smooth ante-carinal space, and raised marginal fold 
of the latter group contrasting strongly with the concentric costation, 
which in Indotrigonia tends to obliterate the distinction between 
area and flank. These characters, together with the marked posterior 
extension and anterior truncation in Pleurotrigonia, justify the 
recognition of two sub-genera which presumably represent divergent 
stocks from a common source. 

The species described below, 7. pongolensis sp. nov., is strikingly 
similar to, and apparently closely allied to, the Hast African 7’. krenkeli 
Lange, which has been compared with T. smeei by both Lange and 
Dietrich, and, indeed, included in Indotrigonia by the latter author. 
The new species differs, however, from the type of Indotrigoma in 
that the costae after the youthful stage do not pass over on to the 
area, which is smooth, and in the highly significant fact that the 
juvenile stage is not characterised by a sculpture of Costate type 


306 Annals of the South African Museum. 


but by a simple series of coarse concentric ribs which pass un- 
interruptedly across the area. The same features are described by 
Lange (1914, p. 231) in the case of T. krenkeli, and I have no doubt 
that these two species are only superficially similar to T. smeez and 
can only be remotely related, and, indeed, may be of quite a different 
origin. I am well aware that certain other species have been in- 
cluded in Indotrigonia or compared with T. smeer that differ from 
that species in important respects. JT. crassa Kitchin (1903, p. 44, 
pl. iv, figs. 4-6; pl. v, figs. 1-3) fails to exhibit the primitive Costate 
ornamentation, though in adult features strikingly ike T. smeei, and 
the failure may be ascribed to loss of the Costate characters at such 
an early stage that they have not been detected, though this is by 
no means certain. T. dietrichi Lange (1914, p. 233, pl. xx, fig. 7) 
has been included in Indotrigona by Dietrich (1933, p. 32, pl. u, 
figs. 38-41), in spite of smaller size, crescentic form, and smooth area, 
on account of the finely developed juvenile Costate stage described 
by the latter. The inclusion of T. crassa and T. dietrichi in Indo- 
trigonia is justified in view of the fact that they have in common 
with T. smeei an essentially concentric flank ornamentation of smooth 
costae and a known or presumed Costate juvenile stage. 

With T. pongolensis and T. krenkeli, however, the position is 
somewhat different. The fact that in these species an initial con- 
centrically ribbed area soon passes into a smooth condition, which 
is maintained to full maturity, indicates descent from a type which 
was concentrically ribbed in the adult stage. It might be the case 
that we have in these two species a development from forms like 
T. smeei, as has been assumed by Dietrich (loc. cit.), but there are 
several considerations which have led me to believe that they are of 
quite different origin, and the remarkable similarity between them 
and T. smeei would therefore be a striking example of convergence 
in evolution. The early loss of the concentric ribbing of the area 
would point to a remoteness in the supposed relationship between 
these species and J. smeez, which it is difficult to reconcile with the 
fact that T. smeei is known to persist into the Zone of T. schwarza 
(Lange, 1914, p. 228; Kitchin, 1929, p. 195), from which T. krenkela 
was derived (Dietrich, 1933, pp. 31-77). The failure of the costae 
to maintain a course strictly parallel to the ventral border—they 
bend upwards when traced posteriorly—indicates an evolutionary 
trend which is not in evidence in T. smeev. On the other hand, a 
primitively concentric ornamentation, with ribs passing from the 
flank across the area, is characteristic of the neanic stage of other 


Lower Cretaceous Lamellibranchia from Northern Zululand. 357 


groups of Trigoniae, e.g. Lotrigona (T. vau, T. v-scripta), Megatrigonia 
(I. conocardirformis) whose relationship to the Costatae has not been 
suggested. A remote kinship with these groups is possible, in spite 
of the very different adult features. 

T. pongolensis and T. krenkelt have several features in common 
with the species included in the T. excentrica Group, for which the 
sub-generic term Rutitrigonia appears to be available. The hetero- 
geneous character of the group of the Glabrae (=Laeves) of Agassiz 
was realised by Lycett (1872, p. 7) and has been referred to by 
various workers, e.g. Bigot (1892, Mém. sur les Trigonies, Mém. Soc. 
Linn. Norm., xvii), who proposed the sub-divisions Semi-Laeves, 
Gibbosae, and Excentricae, the two latter for sub-groups recognised 
by Lycett. Kitchin (1903, pp. 52, 60-62) has pointed out that the 
sub-division appears to be a natural one. Deecke’s term Laevi- 
trigonia (1925, p. 68), proposed for the Glabrae, should be confined 
to the section Gibbosae, since Dietrich (1933, p. 35) cites T. gibbosa 
Sow. as genotype, this being the first citation of genotype that I 
have been able to discover. The distinction between the typically 
Jurassic Gibbosae and the Cretaceous Excentricae has been drawn 
by the authors cited above and need not be elaborated here. 

The term Rutitrigonia was proposed by van Hoepen (1929, p. 31) 
for a new species of the T. excentrica Group from the Cretaceous of 
Zululand, with the rank of a genus and as the sole representative 
of a sub-family Rutitrigoniinae. The type species, R. peregrina 
van H., is characterised by the following features, which have been 
briefly summarised also by Crickmay (1932, p. 459) :— 

Shell moderate in size, inflated and highest in the anterior half, 
posterior part narrowed and produced. Anterior margin broadly 
convex, merging quite gradually into the long convex ventral margin; 
postero-dorsal margin relatively long, concave, sloping gradually, 
downwards so as to meet the ventral margin in a well-rounded posterior 
extremity. Flank ornamented with numerous rather fine, smooth, 
concentric costae. Area relatively narrow, smooth, slightly convex 
in cross-section, indistinctly demarcated from flank and escutcheon, 
without a longitudinal furrow; the marginal and inner carinae are 
represented only by convexities, the former very indistinct; 
escutcheon rather narrow, concave, smooth. Neanic stage with fine 
concentric ribs which at first persist as far as the postero-dorsal 
margin, bending sharply upwards at the margin of the area. Denti- 
tion relatively delicate, the left central tooth broad. | 

The features exhibited by this species are in broad essentials 


358 Annals of the South African Museum. 


identical with those of 7. excentrica Parkinson, as described by Lycett 
(1872-1879, p. 94), noteworthy being the similarity in outline, the 
smooth concentric costae, the indistinctly limited smooth convex 
area, and the fine concentric ribs which are said to cross the entire 
valve near the umbo, the latter characteristic being displayed in 
one of the accompanying figures (zbid., pl. xx, fig. 6). The applica- 
tion of van Hoepen’s term Rutitrigonia to the Excentricae is therefore 
amply justified, the genotype being, indeed, a fairly typical member 
of the latter group; it is perhaps to be regretted that the type of 
the Linnaean group, which now replaces the long-established Excen- 
tricae, is a new species from a remote South African locality, and 
not the English species which gave its name to the Excentricae and 
which has been very well described by Lycett and others. Van 
Hoepen (1929, p. 33) cites as species of Rutitrigonia, T. laeviuscula 
Lycett and ZT. longa Agassiz, both of which were associated with 
T. excentrica by Lycett (1872-1879, pp. 7, 97). It would appear 
also that the East African Neocomian T. schwarz Miller (Lange, 
1914, p. 231, pl. xix, figs. 4,5; Dietrich, 1933, p. 35, pl. im, figs. 52, 
53) belongs to Rutitrigonia, the concentrically ribbed umbonal region 
having been finely figured by Dietrich, and the flank and area having 
the essential characteristics of the Excentricae; the species was 
placed in the sub-genus Laevitrigonia by Dietrich, but has no affinities 
with the Gibbosae for which that term must be used. The rather 
similar T. janenschi Lange and T. bornhardti Miller from East Africa 
belong to the same group. 

It is not without some hesitation that I associate with Rutitrigonia 
the species T'. pongolensis sp. nov. and T. krenkeli Lange. The 
concentric ribbing of the neanic stage is of such a character in both 
these species as to exclude them from Indotrigonia in spite of adult 
similarities, and Rutitrigonia is the only term available for species 
of Trigona having features at all like those of these two species. 
The association of T. pongolensis with Rutitrigonia seems to be 
justified for the following reasons: the anteriorly rounded and 
posteriorly produced form; the concentric smooth costae confined in 
the adult to the flank; the smooth, moderately wide area without 
carinae; the narrow smooth escutcheon; the neanic ornament of 
concentric ribs which pass on to the area. These features include 
the essential characters of the Excentricae. On the other hand, 
T. pongolensis is distinguished from Rutitrigonia in certain particulars. 
The concentric ribbing in the neanic stage is of a remarkably coarse 
character, and the ribs do not bend acutely at the lower margin of 


Lower Cretaceous Lamellibranchia from Northern Zululand. 359 


the area, as do the very much finer juvenile ribs of R. peregrina (which 
“bend back with a swing to cross the area transversely”’), 7’. schwarzi 
(Dietrich, 1933, pl. i, figs. 52, 53) and other species. The costation 
of the flank is of a somewhat different character, for at an early 
stage the costae assume an obliquity, the most notable feature being 
the posterior bending up of the costae; in Rutitrigonia the costae 
tend to be approximately parallel to the ventral margin throughout, 
though it should be pointed out that it has been observed that the 
costae are in part inclined to the growth lines in several species, 
e.g. T. excentrica (Lycett, 1872-1879, p. 94), T. schwarz (Lange, 
1914, pl. xix, fig. 4), so that this distinction must not be regarded as 
of too great importance. The above-mentioned features, which 
serve to distinguish 7. pongolensis from typical Hxcentricae, do not 
seem to be sufficient to warrant the proposal of a subgeneric name, 
and the species is included provisionally in Rutitrigonia with which 
it has many features in common. T. krenkeli Lange is probably 
closely related, and is accordingly removed from Indotrigonia and also 
placed provisionally in Rutitrigonia. 


Trigoma (Rutitrigonia) pongolensis sp. nov. 
(Plate XUil) fies. 5, 6; Plate XLII, figs. 5-7.) 


Material.—Two specimens collected by Dr. 8. H. Haughton and 
Dr. A. W. Rogers on the Mfongosi tributary of the Pongola River, 
both from Z2. The holotype is an almost complete left valve which 
has been developed to show the teeth. The paratype has the valves 
closed and is somewhat damaged posteriorly, but shows the character 
of the ribbing in the vicinity of the umbones. Both specimens are 
comparatively well preserved, but the removal of matrix from the 
surface has been somewhat difficult and I am not quite certain of 
the exact character of the inner carina and its neighbourhood. 

Description of the Species.—Shell of medium size, considerably longer 
than high, with the umbo at about a fourth of the length from the 
anterior end, highest at the umbo, the posterior part slightly narrowed. 
Umbo moderately prominent, well incurved and distinctly opistho- 
gyrous. Shell evenly and moderately inflated, the greatest inflation 
being in front of the middle. Anterior margin sloping steeply 
downwards from the umbo, at first moderately convex, becoming 
more convex below the middle and then merging gradually into the 
long, gently convex ventral margin. Postero-dorsal margin relatively 
long, straight, sloping backwards at a small angle, meeting the inclined 


360 Annals of the South African Museum. 


and nearly straight posterior margin in a very obtuse angle. Posterior 
margin relatively long, meeting the ventral margin in a rounded 
postero-ventral angle, which at middle growth is very obtuse and 
much rounded, but which becomes progressively more acute with 
continued growth; in middle growth the posterior margin is dis- 
tinctly convex and merges almost gradually into the ventral margin, 
but the posterior margin becomes almost straight at a later stage; at 
about 45 mm. from the umbo the posterior margin is about per- 
pendicular to the ventral margin, but at a later stage these margins 
are inclined at an angle less than 90°. 

In front the flank bends down rather rapidly to the anterior margin, 
so that there is a narrow ill-defined steeply inclined anterior face. 
Area moderately wide, becoming progressively wider towards the 
posterior margin, inclined to the flank at an obtuse angle, separated 
from the flank by a broad convexity of the valve surface, the con- 
vexity becoming progressively wider when traced backwards; towards 
the umbones the area is more steeply inclined. Lower part of the 
area gently convex, upper part gently concave, there being either 
no trace of the longitudinal furrow in the posterior part of the area 
(paratype), or an obscure broad furrow well above the middle (holo- 
type). Hscutcheon elongate, narrow, very indistinctly separated 
from the area by a broad convexity which persists to the postero- 
dorsal angle. 

Juvenile ornamentation of a few extremely coarse, elevated, 
smooth concentric ribs separated by rather narrower interspaces, 
the ribs passing over an obscure marginal angle on to the area, 
bending slightly on doing so. There are about 5 ribs in a distance of 
8 mm. from the umbo. 

Adult ornamentation of coarse, elevated, rounded, smooth more or 
less concentric costae, separated by wide concave interspaces. The 
costae are at first inclined to but later about perpendicular to the 
anterior margin, and sweep round in a broad curve so as to be nearly 
vertical as they approach the margin of the area in the middle part 
of the valve. The costae are slightly inclined to the growth lines at 
the anterior end, and when traced backwards appear to cut across 
the posterior part of the valve. The costae cease more or less abruptly 
at the commencement of the anterior slope, the narrow anterior face 
having only growth lines. Traced backwards the costae at first pass 
on to the lower part of the area to die away gradually thereon; ata 
later stage, 15-35 mm. from the umbo, the costae cease on the marginal 
fold; with maturity the costae fail to reach the fold. The area and 


Lower Cretaceous Lamellibranchia from Northern Zululand. 361 


escutcheon after the youthful stage are ornamented only with growth 
lines. ; 
Hinge of the left valve with a broad triangular central tooth, 
deeply embayed below and furrowed on the summit, strongly grooved 
on the sides. Left anterior tooth nearly vertical, grooved on the 
posterior side. 
Dimensions expressed in millimetres: 


Holotype. Paratype. 
Length ; = 00 602 
Height : seed 48 
Thickness of one valve. 12 13 


Remarks.—Reasons have been given above for referring this species 
to Rutitrigonia. The species appears to be very close to the East 
African T. krenkela Lange (1914, p. 230, pl. xx, fig. 2) which was 
included in Indotrigonia by Dietrich (1933, p. 31). The latter species 
was founded on a single imperfect left valve which was said to come 
from the 7. schwarz Beds in all probability; Dietrich recorded the 
species from the same beds, but has not figured additional examples. 
Lange described a narrow, smooth, perpendicular, anterior face, and 
the earlier ribs are said to traverse the area without interruption; 
the area is said to be smooth for the greater part of its length and 
traversed by a shallow furrow, while the escutcheon is also smooth. 
The figure given shows a medium-sized posteriorly produced valve, 
ornamented with widely spaced strong smooth costae which bend 
upwards on the broad marginal fold of the area; the area is shown 
to be steeply inclined, and the escutcheon must be narrow. These 
features and the figure indicate that T. krenkeli and T. pongolensis 
are very closely allied. The species appear to differ only in minor 
respects, T. krenkeli having a much more steeply inclined area and 
T. pongolensis having the ventral and postero-dorsal margins more 
nearly parallel. 


Sub-genus Pleurotrigonia van Hoepen 1929. 


The sub-genus Pleurotrigonia was proposed by van Hoepen (1929, 
p. 33) with the rank of a genus and as the sole representative of a sub- 
family Pleurotrigoniinae, for the single Zululand species 7. blancken- 
horni Newton (1909, p. 40, pl. v, figs. 1-4). The type species has been 
described by both authors and a brief diagnosis of the group has been 
published by Crickmay (1932, p. 459). 


362 Annals of the South African Museum. 


The sub-genus is characterised by the following features :— 

Shell moderate in size, triangular in outline and not much inflated, 
with the umbo near the anterior end and the posterior produced. 
Anterior margin gently convex, with a narrow, elongate, steep, anterior 
face (lunule) separated from the flank by a well-marked anterior 
carinal fold. Postero-dorsal margin long, more or less straight, 
sloping postero-ventrally to meet the ventral margin in a rounded 
angle. Umbo weakly incurved, not appreciably opisthogyrous. 
Dorsal area smooth, moderately wide, with an obscure longitudinal 
furrow, longitudinally ribbed in the youthful stage only; marginal 
carina sharp and nodose near the umbones, passing posteriorly into 
a conspicuous, broad, rounded fold, relatively straight. Escutcheon 
steep, elongate, very narrow; inner carina sharp near the umbo. 
Flank ornamented with strong, rounded, smooth, concentric costae. 
Dentition relatively massive, the left central tooth narrow and only 
obscurely bifid. 

Newton was uncertain of the relations of the type species, but 
suggested that it was an intermediate form coming between the 
Clavellatae and Costatae. That Pleurotrigonia is a derivative of 
the Costatae, can hardly be doubted on the evidence of better speci- 
mens; the smooth concentric costae, the ante-carinal space, the youth- 
ful ornament of sharp carinae and dorsal longitudinal ribs are proof 
of such a relationship; the sub-genus would appear to be an offshoot 
of the Costatae characterised by the complete disappearance, after 
the neanic stage, of the dorsal ornamentation of longitudinal nodose 
carinae and costellae, the marginal carina remaining only as a broad 
conspicuous fold. 

The typically Jurassic Costatae appear to have given rise to a number 
of divergent types towards the close of that period and in the Neo- 
comian, types having in common a trend towards obliteration of the 
longitudinal ornamentation of the dorsal area, unaccompanied by 
any radical departure from the essentially concentric character of 
the flank sculpture. Derivation from the Costatae is indicated in 
many cases by the presence of longitudinal carinae and costellae, 
often nodose, in the early neanic stage. The sculpture of the flank, 
though remaining essentially concentric, may be variously modified, 
but the costae do not become typically nodose or tuberculate. This 
rather heterogeneous group of costate-derivatives, with concentric 
flank ornament and smooth or concentrically ornamented areas, is 
particularly well represented in the Oomia Beds of Cutch (Kitchin, 
1903, pp. 39-60) and includes the species 7. smeer J. de C. Sowerby, 


Lower Cretaceous Lamellibranchia from Northern Zululand. 363 


the genotype of Indotrigonia Dietrich (1933, p. 30). Pleurotrigonia 
falls in the same broad group, though of post-Neocomian age. Some 
of the Indian species, as well as Sphenotrigonia nov., are related types 
marked in addition by the weakening or even obliteration of the 
concentric costation, giving rise to extraordinarily smooth Trigoniae 
that would formerly have been placed in the heterogeneous group of 
the Glabrae; the tendency to smoothness with maturity is exhibited 
also in Pleurotrigonia. 


Trigoma (Pleurotrigonia) blanckenhorni R. B. Newton. 
(Plate XLII, figs. 1-4.) 


1909. T. blanckenhornt Newton: Trans. Roy. Soc. 8. Afr., vol. i, 
p. 40, pl. v, figs. 1-4. 

1929. Pleurotrigonia blanckenhorna (Newton): van Hoepen, Kryt- 
fauna Soeloeland, Trigoniidae, p. 33, pl. vii, figs. 17-19. 


Material.—F our left valves, collected by Dr. 8. H. Haughton and 
Dr. A. W. Rogers at a locality (Z25) south-east of Mokatini, south 
of the big bend of the Pongola River. Two of these shells are com- 
plete and three moderately well preserved. The early neanic stage 
is not present. 

Remarks.—The species was founded on three right valves described 
by Newton as fragmentary, and van Hoepen’s account was based on 
three imperfectly preserved left valves. The species has been well 
described by these authors in so far as the available material enabled 
them to do so. The specimens before me agree pretty well with the 
descriptions and figures given by both authors, and I have little doubt 
that the same species is represented. The new specimens, however, 
would appear to be more finely preserved than those hitherto known, 
and some additional observations can be recorded. 

The anterior region is described by Newton as follows: ‘Anterior 
margin abrupt, deep, smooth, lamelliform,”’ and the concentric 
costae of the flank “‘proceed to within a short distance of the outer 
margin, where they curve upwards by tapering terminations and 
become merged in the growth lamellae of the shell.’”” Van Hoepen 
refers to ‘‘a long narrow lunule which begins at the umbo and ends 
at the lower end of the anterior margin,’ and the costae “begin at © 
the thickened margin of the lunule or a little in front of it.” Neither 
description is quite adequate and the published figures do not clearly 
convey the true character of the anterior region. In the specimens 


6¢ 


364 Annals of the South African Museum. 


before me the gently convex flank of the valves is separated from 
a narrow elongate anterior face or lunule by an abrupt change of 
slope, the anterior face being nearly perpendicular to the plane of 
the valves and to the surface of the flank. The anterior face forms 
with the flank a sharp anterior carina, which, like the anterior margin, 
is only very gently convex forwards, so that the valve is truncate in 
front. The anterior carina is convex as seen from the front, the 
anterior face being widest in the middle and narrowing gradually 
towards both the umbo and the rounded antero-ventral corner. The 
anterior carina is raised and, in the specimens before me, ornamented 
with a row of conspicuous, large, elevated, rounded, or pointed 
tubercles; the tubercles are more or less transversely oval in section, 
with the longer axes parallel to the lines of growth. In many cases 
each of the costae merges into a tubercle at the anterior carina, but 
one tubercle may correspond to two costae, or one of the costae may 
fail to reach the carina, the corresponding tubercle being absent. 
These tubercles, a very conspicuous feature of the shells in the present 
collection, were not mentioned by Newton and van Hoepen, and are 
barely indicated on the figures, their apparent absence being pre- 
sumably due to the state of preservation of the originals. The 
thickness of the shell substance is so great that the anterior carina 
is not noticeable from within as a fold of the valve; the anterior face 
is strictly speaking only a widely bevelled anterior margin. 

Newton observes that the costae of the flank “proceed from” the 
marginal carina, and the accompanying drawings bear out the state- 
ment to some extent, though it would appear that the earlier costae 
just fail to reach the carina. Van Hoepen states that the costae 
‘“‘do not reach the marginal carina in our left valves.” In the speci- 
mens before me the costae near the umbones, 10-15 mm. from the 
apex, terminate posteriorly in slight swellings, leaving a narrow, con- 
cave ante-carinal space, which is crossed by growth lines quite strongly 
concave towards the ventral margin. Traced ventrally the ante- 
carinal space becomes progressively wider and relatively shallower, 
the costae dying out gradually at some distance from the carina, and 
the growth lines crossing the space become less embayed, while the 
costae themselves tend to pass into broad weak corrugations. 

Newton and van Hoepen were not able to describe the earlier 
part of the area, the latter merely observing that the marginal carina 
is at first sharp. One of the specimens in the collection before me 
shows the following essentially Costate youthful features. The 
marginal carina is sharp, elevated, and very conspicuous for about 


Lower Cretaceous Lamellibranchia from Northern Zululand. 365 


15-18 mm. from the umbo, and at 10-15 mm. is unmistakably 
ornamented with fine transverse nodes. The area is gently and evenly 
concave, at least the upper half being ornamented with longitudinal 
ribs. I have not been able to clean the surface thoroughly for fear 
of causing damage, but at 10-18 mm. the upper half of the area 
has unmistakable rounded longitudinal ribs, 3-4 in number, which 
gradually die away at about 20 mm.; I strongly suspect that the 
lower half of the area is similarly ribbed. The inner carina is raised 
and very conspicuous for about 15 mm., and is ornamented with fine, 
rounded, closely set nodes; at about 10 mm. there is a similar, narrow, 
raised, similarly nodose costella immediately below the inner carina, 
and separated from it by a deep narrow furrow, with a suggestion 
of a similar costella below it; in a perfect specimen it is probable 
that the area would be found to have a number of raised nodose 
costellae, which become smooth and finally die out when traced 
posteriorly at about 20 mm. from the umbo. 

The escutcheon is very narrow and at first very strongly marked 
off from the area by a very abrupt descent of the valve surface, the 
inner carina being very sharp. Traced posteriorly after 15 mm., 
the escutcheon and area gradually merge into one another. At first 
the escutcheon is deeply grooved above the middle, the almost vertical 
lower part being rather sharply distinct from a very narrow shelf-like 
upper margin. Newton states that the escutcheon is insignificant, 
and even in his comparatively large shell only about $ mm. wide. 

The identification of the specimens before me with Newton’s types 
is greatly strengthened by the fact that the former were found 
associated with Haogyra cf. conica (J. Sow.) and Trigonarca ct. 
lagertensis (d’Orb.), two species which occur at the type locality, 
and with Veniella etheridgei Newton, a characteristic Albian species 
from localities farther south. 


Sub-genus Sphenotrigonia sub-gen. nov. 


The new sub-generic term Sphenotrigonia is here introduced for 
a very remarkable new species of Trigonia from the Lower Cretaceous 
of Northern Zululand, having the size and something of the shape 
of a coup-de-poing. The genotype, I. frommurzei sp. nov., is described 
below. The species is characterised by the remarkably long and 
almost perfectly straight postero-dorsal margin, the long and almost 
straight central margin, the excessively produced straight and 
pointed posterior region, the very marked anterior truncation, which 

VOL. XXXI, PART 3. 30 


a 


366 Annals of the South African Museum. 


combination of characters gives the shell a very distinct shape like 
that of a wedge; to these characters is added the complete dis- 
appearance of costae and dorsal carinae after the neanic stage. The 
hinge is in all essentials that of a Trigonia. 

The unusual form of the species led to the assumption in the field, 
and until the development of the hinge in suitable specimens had 
been completed, that a second South African species of the aberrant 
genus Seebachia had been discovered, an interesting comparison, 
since the genotype of the latter has been placed in the Trigoniidae 
by certain authors. The general resemblance between T. frommurzei 
and Seebachia bronni is indeed very remarkable, in spite of the 
presence of a number of external features which serve to distinguish 
them, and a very different dentition having only superficial points 
in common with Trigonia. Since Seebachia bronni, until recently the 
only known species, has not been described since 1881, and is rare 
in collections, and has been quoted as a member of the Trigoniidae 
as recently as 1925, it is perhaps opportune to point out the characters 
which serve to exclude Seebachia from that family.* 


* The Systematic Position of Seebachia bronni (Krauss). The genus Seebachia 
was proposed by Neumayr (1881, p. 274) for a species from the Upper Valanginian 
Sundays River Beds in South Africa originally described as an Astarte by Krauss 
(1850, p. 449, pl. xlviii, fig. 1, a-e), A. bronni. Krauss regarded the species as 
nearest to Astarte but probably the representative of a new genus “between” 
Astarte and Trigonia. Neumayr saw in the peculiar elongate form and grooved 
teeth a great distinction from Astarte, and while admitting that the character of 
the teeth recalled T’rigonia in certain respects, expressed the opinion that the 
genus was not closely related to the latter. 

Both Krauss and Neumayr gave excellent figures showing the internal and 
external features, Krauss of both valves, Neumayr of a right valve. These 
figures, together with the descriptions that accompany them, are sufficient to 
indicate whether the species has or has not affinities with Trigonia. It may be 
pointed out that there appear to be very few specimens of Seebachia bronni in 
existing collections; in addition to the two specimens known to Krauss, and the 
sight valve figured by Neumayr, I have been able to trace only three specimens 
of the species; one in the Albany Museum in Grahamstown (A.M. 2418), 
a finely preserved specimen with the valves closed from the Sundays River, one 
in the South African Museum in Cape Town (S.A.M. 4676), from Picnic Bush 
on the Zwartkops River, a left valve which with much labour was developed to 
show the dentition; and a specimen in the Geology Department of the University 
of Cape Town. 

That Seebachia bronni does not belong to the Trigoniide is shown clearly 
enough by the character of the hinge, and is borne out by other characteristics 
of the shell. In the left valve there are two teeth only, the larger and posterior 
of the two, which is in the position of the bifid tooth of a Trigonia, is oblique, 
elevated, narrow, and entire. In the right valve there is a large elevated tooth 


Lower Cretaceous Lamellibranchia from Northern Zululand. 367 


The sub-genus Sphenotrigonia sub-gen. nov. (Type: 2. frommurzei 
Sp. nov.) 

Diagnosis.—Shell large, massive, elongately triangular in outline, 
highest and most inflated close to the anterior end, and tapering 
very gradually by diminution in height and thickness to a posterior 
point. Anterior end markedly truncate, with a more or less vertical 
carinal fold and anterior margin, and a flattened anterior face. 
Posterior very much produced, the postero-dorsal and ventral margins 
very long and straight. Primitive ornamentation of oblique costae 


reminiscent of the right anterior tooth of the latter genus and a narrow posterior 
tooth. Affinities with T’rigonia were presumably suggested by the circumstance 
that both sides of the large right tooth and the opposed sides of the left teeth 
are strongly grooved, and it might be held that Seebachia is an aberrant Trigoniid 
in which (i) the posterior tooth of the left valve has disappeared and in which 
(ii) the left central tooth has become narrower so as to loose its bifid character, 
with a loss of the grooves on its posterior side. The remarkable stability shown 
by the Trigonia dentition throughout its Jurassic-Cretaceous existence should 
put us on our guard against such an interpretation, and it has been pointed out 
that transversely striate teeth are not the monopoly of that genus (Crickmay, 
1932, p. 453). 

The hinge apparatus is further distinguished from that of T'rigonia in a very 
important way, which, in conjunction with the above features, renders any reference 
to the Trigoniidae out of the question. In S. bronni the teeth are situated on a 
well-developed shelf-like hinge-plate, which has no counterpart in the Trigoniae; 
the difference is particularly evident in the right valve, where in S. bronni the 
large central tooth arises from the hinge-plate and receives no support directly 
from the floor of the valve cavity as is the case with the large anterior tooth of 
the Trigoniae. In addition S. bronni is characterised by a deeply concave lunule 
and denticulations within the ventral border, features which are not normal to 
the Trigoniae. Seebachia has, in recognition of these features, been placed in the 
Astartidae by various authors, e.g. in Zittel (1913, p. 473), by that authority on 
the Trigoniae the late F. L. Kitchin (1929, p. 194), and by Dietrich (1933, p. 42), 
who has described a second species from the 7’. smeei-Beds in Tanganyika. 

Several authors have followed Stoliczka in including S. bronni in the Trigoniidae. 
Stoliczka (1871, p. 312), acquainted only with the paper by Krauss (1850), listed 
the species in that family as a species of Remondia, a genus placed by its author 
Gabb (1869, p. 270) in the Trigoniidae, but which has been clearly shown by 
Stanton (1897) to be in no way related to that family, having an essentially 
heterodont dentition with cardinals and laterals. S. bronni has nothing in 
common with Remondia, a genus of smaller shells with a different external appear- 
ance and quite different dentition, and it is difficult to understand Stoliczka’s 
reference. Yet Cossmann (1912, pp. 59-84) in his work on the evolution of the 
Trigoniae not only accepted Remondia as an aberrant offshoot of the Trigonia 
stock, but quoted S. bronni as a species of that genus, and Deecke (1925, pp. 58-59), 
in his catalogue of the Trigoniae, included both Remondia and Seebachia, which 
have been clearly shown by authors to belong elsewhere. 


368 Annals of the South African Museum. 


and dorsal carinae restricted to the neanic stage; costae slightly 
oblique, smooth (?), terminating posteriorly in a row of tubercles 
in advance of the marginal carina; marginal and inner carinae 
delicately beaded; dorsal area and escutcheon distinctly limited 
only in the neanic stage, the area with smooth (?) longitudinal 
costellae above the median sulcus and the escutcheon quite smooth. 
The greater part of the shell devoid of costae or dorsal carinae, 
ornamented only with growth lines, and broad concentric undulations 
towards the anterior end. Dorsal area beyond the neanic stage 
relatively narrow, passing imperceptibly into the flank, without 
carinae, ornamented only with growth lines, without escutcheon. 
Hinge massive, the right posterior tooth very long, and grooved 
only in the proximal half, with a corresponding very long left posterior 
socket. 

There are not many Trigonie with which this well-marked type 
can be compared. There is a resemblance in certain respects to 
Pacitrigonia Marwick (1932) founded on an upper Senonian New 
Zealand species, which is ornamented only with growth lines beyond 
the neanic stage, and is a large posteriorly produced shell, with 
oblique costae nearly parallel to the posterior ridge or marginal 
carina only in the neanic stage. The New Zealand species, however, 
is compressed and does not possess the extraordinarily attenuated 
posterior and strong anterior carinal fold so characteristic of Spheno- 
trigonia. The area of Pacitrigonia is described as smooth and the 
escutcheon and inner carina are said to be absent, but the neanic 
stage of these appears to be unknown. 

There is a considerable resemblance in general features between 
Sphenotrigonia and Pleurotrigonia van Hoepen. In each case the 
type species is characterised by marked anterior truncation and 
posterior elongation, a juvenile ornamentation of Costate character 
with non-nodose ribs, beaded dorsal carinae and ante-carinal sulcus, 
a relatively narrow area and narrow escutcheon. In Sphenotrigonia, 
however, the costation is essentially a juvenile feature, and the area 
becomes indistinct, whereas in Pleurotrigonia the costation, ante- 
carinal space, and broadly rounded carina are maintained until the 
adult stage, though obsolescent with full maturity. The excessively 
elongate and pointed posterior, very narrow and indistinct area, 
and smooth surface of Sphenotrigonia are peculiar features which 
appear to justify the proposal of a new sub-generic term for 7. from- 
MUrZer. 

The ornamentation of the neanic stage in Sphenotrigonia indicates 


Lower Cretaceous Lamellibranchia from Northern Zululand. 369 


derivation from the Jurassic Costatae, shown by the beaded dorsal 
carinae, the longitudinal costellae at least on the upper part of the 
area, and the oblique smooth (?) costae which appear to be pseudo- 
concentric within a few millimetres of the umbo. The sub-genus is 
no doubt related to the heterogeneous assemblage of species, placed 
by Kitchin (1903) in the convenient group of Costate-derivatives, 
that are linked together by the possession of Costate neanic features, 
concentric adult ornamentation, and smooth areas. The marked 
anterior truncation and extraordinarily pointed posterior are quite 
unique, however. In some of the Indian species, as in Sphenotrigonia, 
the trend towards obliteration of longitudinal dorsal ornament has 
been accompanied by a trend towards weakening or even obliteration 
of the concentric costation of the flank, producing extraordinarily 
smooth Trigoniae which would formerly have been included in or 
associated with the admittedly very heterogeneous assemblage of more 
or less smooth species known as the Glabrae. For the Glabrae, 
Deecke (1925, p. 68) has proposed the sub-genus Laevitrigonia, the 
type of which by subsequent designation (Dietrich, 1933, p. 35) 1s 
T. gibbosa Sow., a Jurassic species of a very different ancestry to the 
smooth types here being discussed. Sphenotrigonia has very little 
in common with and is not related to Laevitrigonia. 

Kitchin (1908, p. 101) has remarked on the tendency to posterior 
elongation, and loss of sculpture on the adult area, shown by several 
unrelated types in the Uitenhage and Oomia beds. In no Trigonia 
is the extension of the siphonal end as extraordinary as in Spheno- 
trigonia, and it is rather curious that in the Indo-African Lower 
Cretaceous the same tendency has produced a similar excessively 
elongated shell in an unrelated family, the Astartid genus Seebachia. 


Trigoma (Sphenotrigonia) frommurzer sp. nov. 
(Plate XLV, figs. 1-3; Plate XLVI, figs. 1-4; Plate XLVII, figs. 1-4.) 


Material.—Nearly 60 specimens of this very remarkable species 
were available for study. The majority of these are more or less 
complete individuals with the valves closed and the surface roughened, 
pitted, or otherwise damaged, the specimens having been collected at 
the surface after their release from the parent rock by the agents 
of weathering. A number, however, are comparatively well preserved, 
and several specimens show very finely the internal characters of the 


two valves. 
The species was collected by Mr. J. 8S. Hutt and by Mr. H. F. 


370 Annals of the South African Museum. 


Frommurze along the Mfongosi tributary of the Pongola River 
(T.M. 1667, 1669, 1672, 1674-1677, 1680, 1681, T683)eaag 
S.A.M. 10585-10592), the horizons not being noted. In 1933 
the locality was visited by Dr. 8. H. Haughton and Dr. A. W. Rogers, 
who collected about 40 specimens along the Mfongosi stream at 
several horizons. The species appears to be characteristic of several 
of the localities, and was recorded from each of the localities Z6—Z13. 
The number of specimens collected from each is indicated in brackets: 
Z6 (7), ZT (13), Z8 (10), Z9 (2), Zi@ (2), ZL (1), 712 (1) eae 

The species was also found in L3 (3 specimens) on the Lombagwenya 
stream to the north. 

Holotype.—The most complete specimen available is taken as 
holotype of the species, in spite of the fact that the internal characters 
are not shown, and although the specimen is from a horizon on the 
Mfongosi stream not precisely known. It so happens that all the 
specimens showing the hinge are unsatisfactory in that they are 
either small or are full-sized specimens very incomplete in the posterior 
part. The holotype (S. Afr. Mus. 10586) was collected by H. F. 
Frommurze, and is quite complete even to the tip of the posterior; 
the valves are tightly closed and in a fair state of preservation, 
though the flanks are rather badly pitted. 

Description of the Species. (1) External Features.—Shell large, 
massive, equivalve, extremely inequilateral, elongately triangular in 
outline, about twice as long as high, anteriorly very markedly truncate, 
with the greatest height and thickness very close to the anterior, 
tapering very gradually from the anterior to a posterior point. 
Umbones small, situated very close to the anterior end, highly 
incurved and contiguous or nearly so. 

Anterior margin moderately long, approximately vertical, very 
gently and evenly convex from the umbones to the well-rounded 
antero-ventral angle. Ventral margin long, about twice as long as 
the anterior margin, moderately convex in the anterior half, becoming 
almost straight or even slightly concave in the posterior half. Postero- 
dorsal margin long, about as long as the ventral margin, remarkably 
straight throughout its length. Postero-dorsal and ventral margins 
meeting in a rounded point, the two margins forming an angle of 
about 30-40°. Postero-dorsal and anterior margins forming an angle 
of about 70°. 

A strong carinal fold runs almost vertically downwards from the 
umbo to the antero-ventral corner. Carinal fold almost straight, 
slightly convex towards the anterior, clearly dividing the external 


Lower Cretaceous Lamellibranchia from Northern Zululand. 371 


surface of the valve into two regions, which meet at the carinal fold 
at an angle not much greater than a right angle (about 100°), a narrow 
anterior region being separated from the broad flank of the valve. 
Anterior region flattened or gently convex. When closed the two 
valves are cordate to narrowly cordate in anterior aspect. Carinal 
fold somewhat sharply to gently rounded. 

Greatest convexity of the shell close to or a little posterior to the 
~ carinal fold, the thickness decreasing quite gradually from the neigh- 
bourhood of the carinal fold to the posterior extremity. When 
closed the two valves are narrowly wedge-shaped in dorsal, ventral, 
or lateral aspect. Valves often slightly or very slightly concave in 
the posterior part, with a very broad and very shallow sulcus running 
obliquely downwards, corresponding to a slight embayment of the 
posterior half of the ventral margin. The sulcus and embayment 
are scarcely discernible in the holotype and other specimens. Broad 
flank of the valves flattened and only shghtly convex in the middle part. 

For the greater part of the length of the adult shell the flank passes 
over to the straight postero-dorsal margin in a broad, rounded 
convexity devoid of carinae or costae, interrupted only by growth 
lines and by a persistent, narrow, weak, longitudinal furrow towards 
the upper side. For a distance of about 20 mm. from the umbo 
there are two posteriorly directed, finely beaded carinae, demarcating 
the dorsal area and escutcheon. The marginal carina is narrow, 
raised, rib-like, and ornamented with conspicuous rounded tubercles 
for at least 15 mm. from the umbo, giving a beaded appearance; 
the marginal carina curves obliquely backwards, gently concave 
towards the dorsal area, for about 20 mm. from the umbo, and passes 
gradually into a broader fold. For almost the entire length of the 
shell the area is of constant width and relatively narrow, and is not 
clearly demarcated from the flank of the shell, except that the broad, 
rounded convexity has its very much rounded crest,.across which 
the growth lines bend round rapidly, in line with the marginal carina 
near the umbones. Near the umbones the area is relatively broad 
and gently concave. The area is divided longitudinally by a distinct 
groove; near the umbones the groove divides the area into two nearly 
equal flattened or slightly convex bends; traced posteriorly the upper 
and lower limits of the area are ill-defined, but the longitudinal groove 
persists for the greater length of the shell, curving very slightly so 
as to meet the postero-dorsal margin near the posterior extremity. 
Near the umbones there is a distinct escutcheon, elongate, narrower 
than the area, separated from the area by an inner carina similar 


372 Annals of the South African Museum. 


in character to and continued posteriorly for the same distance as 
the marginal carina; the inner carina 1s narrow, sharp, and beaded, 
and above it the escutcheon is abruptly excavated, concave, and 
quite smooth. The ornamentation of the area in the vicinity of the 
umbones is almost unknown; in the holotype, however, there are 
quite unmistakably two longitudinal costellae between the furrow 
and the inner carina. Traced posteriorly the inner carina passes 
into a broad fold, and for the greater length of the shell no distinction 
between area and escutcheon is possible, though there is a slight 
flattening at the valve margin. 

There is a very distinct though only slightly excavated lunule, re- 
presented by a narrowly cordate shallow depression near the umbones 
on the anterior face of the valves, demarcated from the remaining 
part of the anterior region by a change in slope; in adult shells the 
lunule is about 20 mm. high and 13 mm. in width across the two 
valves. 

The shell material is remarkably thick, 4-8 mm. over the greater 
part of the valves in the adult and greatly thickened to support the 
teeth. 

(2) Internal Features.—Hinge apparatus very strongly developed 
and situated close to the anterior end, the teeth stout, elevated, and 
strongly grooved laterally. 

In the left valve there is a strong central bifid tooth separated by 
an anterior and a posterior socket from an anterior and a posterior 
tooth respectively. Central tooth directed obliquely backwards, 
elevated well beyond the hinge margin, elongate-triangular in shape 
and shallowly concave along the summit, with the vertical sides 
diverging and the lower end conspicuously embayed; sides of the 
tooth approximately vertical and strongly grooved, with 12-15 
grooves on the posterior side and 8-10 grooves on the anterior side. 
Anterior socket elongate, very deep, but narrower than the central 
tooth, with the lower end closed in by a raised wall joining the bases 
of the central and anterior teeth. Anterior tooth raised, the posterior 
side nearly vertical and strongly grooved with 8-10 grooves, the 
anterior side convex and smooth. Anterior socket, together with the 
anterior tooth and the anterior fork of the central tooth, supported 
by a broad convex internal rib, which is continued ventrally for a 
short distance so as to separate the anterior adductor pit from the 
valve cavity. Posterior socket moderately deep, rounded, very 
elongate, up to 40-45 mm. long in the adult, directed posteriorly 
parallel to the hinge margin, bounded ventrally by an elevated sharp- 


Lower Cretaceous Lamellibranchia from Northern Zululand. 373 


erested ridge, which is the prolongation of the posterior fork of the 
central tooth. Posterior tooth confluent with the nymph, narrow 
and elongate, its crest appearing as a very narrow ridge against the 
summit of the nymph, strongly grooved on the ventral side up to 
about 25 mm. from the umbo. Nymph relatively narrow and with 
the very narrow ligament groove continued for about 30 mm. from 
the umbo in the adult. 

In the raght valve there is a very elevated anterior tooth and a very 
elongate narrow posterior tooth. Anterior tooth large, relatively 
narrow, directed ventrally but a little obliquely, very elevated, 
raised well above the hinge margin, up to 12 mm. high in the adult, 
the sides vertical and diverging at a small angle, both sides strongly 
grooved with 8-10 grooves. Anterior tooth supported by a broad 
convex internal rib, which is continued ventrally for a short distance 
so as to separate the anterior adductor pit from the valve cavity. 
Anterior to the anterior tooth is a relatively narrow, moderately 
deep and rather short anterior socket, the anterior side of which is 
concave and smooth. Between the anterior and posterior teeth is 
a broad triangular gap in which there is, against the base of the 
anterior tooth, a narrow shelf formed by a continuation of the 
supporting internal rib. Posterior tooth very long, up to 40-50 mm. 
in the adult, narrow, sharp-crested, situated close to and parallel to 
the hinge margin, with the ventral side free to the floor of the valve 
cavity, and the dorsal side free only along the summit; end nearest 
the umbo overhanging and strongly grooved from the summit to 
the floor of the valve cavity, with corresponding short but strong 
grooves on the dorsal side up to about 25 mm. from the umbones. 
Posterior socket reduced to a very narrow and shallow furrow about 
25 mm. long separating the elongate posterior tooth from the narrow 
nymph, its dorsal margin smooth, its ventral margin being the grooved 
summit of the dorsal side of the posterior tooth. 

The adductor pits are subequal, close to the valve margins and very 
deep. Anterior adductor pits oval, extremely deep, situated so close 
to the anterior margins that the shell is quite thin on their anterior 
side; pits separated from the valve cavity by the internal supporting 
rib and the lower part of the anterior tooth. Posterior adductor pits 
rounded and deep; between each and the posterior end of the hinge 
is a small accessory pit. Pallial line strongly impressed, simple, at 
some distance from the ventral margin in the anterior half. Ventral 
margin smooth within, with a broad, flattened, marginal band along 
which the valves meet when closed. 


HT 


374 Annals of the South African Museum. 


(3) Ornamentation.—The external sculpture consists almost en- 
tirely of concentric growth lines and concentric undulations, the 
valves consequently appearing to be remarkably smooth for a 
Trigonia. Costae, as well as the dorsal carinae, are strictly confined 
to the neanic stage, and do not appear beyond about 20 mm. from 
the umbo. Flanks of the valves with numerous relatively fine 
growth lines which follow the curvature of the ventral margin and 
towards the umbo pass obliquely across the costae. The growth 
lines bend somewhat rapidly across the anterior carinal fold so as 
to follow straight courses across the anterior face and the lunular 
depression, inclined very obliquely to the anterior margin. Towards 
the postero-dorsal margin the growth lines bend round in a broad 
curve over the dorsal convexity, which represents the lower ill- 
defined boundary of the dorsal area, and are inclined very obliquely 
to the margin. In the anterior half the surface of the flank is 
characterised by a series of about 10 broad concentric undulations, 
with broad moderately convex crests and concave or broadly 
V-shaped interspaces, 6-8 mm. apart, representing a more or less 
regular growth periodicity. These concentric undulations appear 
only after the neanic stage and may not be maintained towards the 
ventral margin. The undulations vary in strength from shell to 
shell, being strongly developed in some, while others are apparently 
marked by a more or less even surface. The undulations are strongest 
near the anterior carinal fold and become progressively weaker when 
traced posteriorly, failing to reach the smooth postero-dorsal con- 
vexity; they reach the carinal fold and cross over it on to the 
anterior face, but then very rapidly disappear, the greater part of 
the anterior face being crossed only by oblique growth lines. 

Near the umbo, in the neanic stage, is a series of regular, relatively 
elevated, rounded, slightly oblique costae, about 10-12 in number, 
which are almost parallel to the marginal carina bounding the dorsal 
area and separated from each other by rounded interspaces of about 
the same width as themselves. The costae persist to about 20 mm. 
from the umbo. The costae commence at the anterior carinal fold 
where they end abruptly, and are a little swollen in some specimens; 
the costae cease posteriorly in an oblique row of conspicuous tubercles, 
leaving a rapidly widening smooth oblique band between the costate 
region and the beaded marginal carina. The costae appear to have 
been devoid of tubercles or granules along their summits, but this 
absence may be due to wear. 

The beaded marginal and inner carinae have been described above 


Lower Cretaceous Lamellibranchia from Northern Zululand. 375 


in connection with area and escutcheon. The beaded character is 
apparent only in young specimens, the tubercles having been worn 
away in the adult shells. 

(4) Dimensions (expressed in millimetres) : 


S.A.M. No. . . 10586 10587 10588 10590 
Length . : 5 Sti oy x 
Height . ! : : 69 79 65 73 
Thickness of one valve . 23°5 27 28-5 20 
Postero-dorsal margin . 128 a he 


Remarks.—The degree of inflation of the valves varies very con- 
siderably, the holotype being rather narrowly cordate in anterior 
aspect, while other specimens may be more broadly cordate. The 
ventral margin in the holotype is straight and there is little sign of 
the broad shallow posterior sulcus; in other examples the sulcus is 
quite evident and gives rise to a conspicuous gentle embayment of 
the ventral margin. These variations were not found to have any 
zonal significance, but this may be due to insufficiency of material. 


Sub-genus Scabrotrigonia Deecke 1925. 
Trigoma (Scabrotrigonia) cricki R. B. Newton. 


1909. Trigonia cricki Newt.: Trans. Roy. Soc. 8. Afr., vol. i, p. 38, 
pl. v, figs. 10-15. 

1929. Ptilotrigonia cricki Newt.: van Hoepen, Krytf. Soeloeland, 
poe, pl vietio. 7; pl: viz figs. 1-4. 

A preliminary examination of the Trigoniae of the section Scabrae, 
which are common throughout the succession and apparently repre- 
sented by several species, shows that J. cricki is represented by a 
fine set of specimens from Z19 (20) on the Pongola River below the 
junction with the Mfongosi stream. The type is from the Albian 
(du Toit, 1926, p. 318) of the Manuan Creek and van Hoepen has 
recorded the species from the Umsinene River, Umsinene Beds, 
identified by him as Albian (1926, p. 222). Dr. 8. H. Haughton also 
recorded the species at locality Z20. 


Trigonia (Scabrotrigonia) ct. kraussi Kitchin. 


1908. 7. kraussi Kitchin: Ann. S. Afr. Mus., vol. vu, p. 95, pl. 111, 
fig, 2. 

Large Scabrae comparable with the Uitenhage species T. kraussi, 

and with 7. salebrosa (van Hoepen) (1929, p. 20, pl. v, figs. 1-6) from 


376 Annals of the South African Museum. 


the base of the succession on the Umsinene River, are not common, 
but Mr. J. 8. Hutt collected two specimens (T.M. 1768, 1769) 
on the Mfongosi stream at an unrecorded horizon, and Dr. S. H. 
Haughton and Dr. A. W. Rogers found a single specimen at Z8, the 
three specimens apparently belonging to a new species allied to 
T. krausst. These will be described in a subsequent communication. 


Trigonia (Scabrotrigonia) spp. 

Several species are probably represented among the numerous 
smaller Scabrae in the collection. An apparently undescribed 
species with exceptionally coarse tubercles is abundant at Z3 (17) 
and Z4 (30) on the Mfongosi, the material being in excellent condition. 
The Scabrae will be dealt with in a separate paper. 


Famity PECTENIDAE. 
Genus NEITHEA Drouet 1825. 


Neithea quadricostata (J. Sowerby). 


1903. Pecten (Neithea) quadricostatus J. Sow.: Woods, Cret. Lam. 
England, vol. i, p. 210, pl. xl, figs. 6, 7. 

1909. Neithea quadricostata (Sow.): R. B. Newton, Trans. Roy. 
Soc. 8. Afr., vol. i, p. 55, pl. u, figs. 18-21. 

1930. Newthea quadricostata (Sow.): Rennie, Ann. 8. Afr. Mus., 
vol. xxvii, p. 241, pl. xxx, figs. 10-12. 


This well-known Albian and Cenomanian species has been figured 
from the Manuan Creek in Zululand by Newton and Rennie, and 
has been recorded (Rennie, loc. cit.) from the Pongola River. The 
present collections include a very fine series of about 50 specimens 
from Z19 on the Pongola River below its junction with the Mfongosi 
stream, as well as an incomplete juvenile convex valve from Z9 
which apparently represents the same species, and 2 good specimens 
from Z20. From L5 (Msinyene Pan) comes a juvenile convex valve 
of a Newthea which may be this species. 


Neithea quinquecostata (J. Sowerby). 
1903. Pecten (Neithea) quinquecostatus J. Sow.: Woods, Cret. Lam. 
England, vol. i, p. 202, figs. on pls. xxix, xl. 
1930. Neithea quinquecostata (J. Sow.): Rennie, Ann. 8. Afr. Mus., 
vol. xxvii, p. 179, pl. xvau, figs: 1-5 (celerenceswim 
S. African records). 


Lower Cretaceous Lamellibranchia from Northern Zululand. 377 


This species is of common occurrence in the Umzamba Beds 
(Campanian) of Pondoland and has been recorded from a number of 
localities in Zululand. In Europe it ranges from the Aptian to the 
Senonian. A single but quite typical convex valve was collected 
from Z13 on the Mfongosi, from an horizon certainly not later than 
Cenomanian. 


Famity PHOLADOMYIDAE. 
Genus PHoLapomya G. B. Sowerby 1824. 
Pholadomya? luynesi Lartet. 
(Plate XLITI, fig. 4.) 


1877. P. luynest Lartet: Géol. Mer. Morte, p. 125, pl. xi, figs. 7, 8. 
1909. Ke be R. B. Newton, Trans. ae SOC, Sp “ute. 
WOE pes ple, Mes. lee2: 


Newton figured a shell from the Manuan Creek in Zululand and 
referred it to this species, which is said to occur in the Cenomanian 
of Syria, though the type is from a higher horizon. There is a single 
well-preserved specimen from the Pongola River below its junction 
with the Mfongosi stream, from Z19, which agrees with the shell 
described by Newton and must be referred to the same species. In 
the collection of the Transvaal Museum there are further excellent 
examples of the species (T.M. 1235, 1273, 1274, 1301) in an Albian 
fauna from Catuane in Portuguese Hast Africa. 

The species has been retained in Pholadomya by Blanckenhorn and 
Newton in spite of the absence of radial ribs, but might be better 
placed in Homomya or even in Pleuromya. 


Pholadomya vignesr Lartet. 


1877. P. vignesi Lartet: Géol. Mer. Morte, p. 126, pl. xi, fig. 9. 
1.908). nt % emend. Blanckenhorn: R. B. Newton, 
Trans. Roy. Soc. 8. Afr., vol. i, p. 79, pl. vi, figs. 3-6. 


Newton has described and figured examples of this species from 
the Manuan Creek and has remarked on its comparative abundance 
at that Zululand locality. The present collection includes a number 
of finely preserved and very typical specimens, whose identity with 
the shells described by Newton is not open to doubt. These were 
collected as follows: on the Lombagwenya stream, L7 (12); in the 
neighbourhood of the Mfongosi stream, Z16 (7), Z19 (10), Z20 (1), 
Z21 (1). There are finely preserved examples of this species in the 


378 Annals of the South African Museum. 


Albian fauna from Catuane, Portuguese East Africa, in the collec- 
tion of the Transvaal. Museum (T.M. 1275, 1303, 1307-9). In 
Palestine, Algeria, etc., the species occurs in the Cenomanian, while 
at the Manuan Creek the species is said to be from the Albian (du Toit, 
1926, p. 318). 


Genus Goniomya Agassiz 1842. 


Goniomya sp. 


The genus is represented by rather fragmentary specimens probably 
belonging to the same species as Goniomya (species 1) figured by 
Newton (1909, p. 81, pl. vi, figs. 18, 19) from Manuan Creek in Zulu- 
land. These were collected as follows: neighbourhood of the 
Mfongosi stream, Z16 (1, large), Z21 (1). A rather similar form is 
present at Catuane (T.M. 1137, 1289), but the poorness of the 
Zululand material makes comparisons difficult. 


Famity CYPRINIDAE. 
Genus VENIELLA Stoliczka 1871. 
Veniella etheridge: R. B. Newton. 


1909. V. etheridger, Newt.: Trans. Roy. Soc. 8S. Afr., vol. 1, p. 69, 
pl. ai ties OPO 

1930. V. etheridgei, Newt. : Rennie, Ann. 8. Afr. Mus., vol. xxvii, 
p. 242, pl. xxxi, figs. 1-3. 


This massive and very distinct species is represented in the present 
collections by a number of specimens, the majority with the valves 
closed and fairly well preserved. The species was described by 
Newton from the Manuan Creek in Zululand, and is said by du Toit 
(1926, p. 318) to be from the Albian. A fine left valve was figured 
by Rennie from the same locality, and the occurrence of the species 
with Neithea quadricostata on the Pongola River was recorded. The 
species has been recorded from: on the Lombagwenya stream, L3 
(12%, juvenile specimen, poorly preserved); neighbourhood of the 
Mfongosi stream, Z16 (3, small or broken), Z19 (19, fine typical 
specimens), Z20 (1); south of Mokatini, Z25 (5). 


Lower Cretaceous Lamellibranchia from Northern Zululand. 379 


Famity CARDIIDAE. 
Genus ProrocarpiA Beyrich 1845. 
Protocardia ct. sphaeroidea (Forbes). 
(Plate XLVIII, figs. 1-6; Plate XLIX, fig. 9.) 


1908. Woods: Cret. Lam. England, vol. ii, p. 195, pl. xxxi, figs. 2 a, 
YAO SO oD. 


Material.—The genus is represented by about twenty fairly well 
preserved specimens of a large species, from exposures on the Mfongosi 
stream and on the Pongola River. Mr. H. F. Frommurze and 
Mr. J. 8S. Hutt collected five specimens on the Mfongosi (S8.A.M. 
NOTSS hO7S9; M1790; 1796, 1894). Dr. 8. H. Haughton and 
Dr. A. W. Rogers collected three specimens on the Mfongosi, one 
specimen from each of the localities Z11, Z12, Z13; and ten specimens 
from Z19 on the Pongola River below its junction with the Mfongosi. 

Remarks.—These large shells are very similar to the English Aptian 
P. sphaeroidea (Forbes), which has been well described and figured by 
Woods (loc. cit.). The type is from the Perna-Bed (lower Aptian) of 
the Isle of Wight. The English species has been described by Woods 
as follows, and the description applies equally well to the Zululand 
specimens: “Shell stout, large, much inflated, higher than long, 
slightly inequilateral. Anterior and ventral margins rounded. 
Posterior margin truncated, forming angles with the postero-dorsal 
and ventral margins. Umbones prominent, with a small forward 
curvature, and an inconspicuous carina extending to the postero- 
ventral angle and limiting the flattened postero-dorsal area. Orna- 
mentation consists of regular, broad, flat concentric ribs separated 
by narrow grooves. On the posterior area strong growth ridges are 
present.” 

Woods figures only two specimens, and I have found it difficult to 
make a satisfactory discrimination between these and the Zululand 
set. There is also a considerable resemblance to P. rothpletz: Krenkel 
described recently by Dietrich (1933, p. 51, taf. vi, figs. 89-91) from 
the Trigonia schwarzi-Beds in the Tendaguru Formation of Tangan- 
yika. The latter species is said by Dietrich to be very closely allied 
to P. sphaeroidea, but to be distinguished by slightly narrower ribs 
and less pointed umbo. From the measurements given by Woods 
and Dietrich for the two species, it would appear that the Hast African 
and English shells cannot be separated by reason of any conspicuous 


380 Annals of the South African Museum. 


difference in size or proportions. The ratio of length : height for 
P. sphaeroidea is from 1: 1-02 to 1: 1-26, the corresponding figures 
for P. rothpletza being 1: 1-03 to 1: 1-06, which is within the range 
given for the former species. 

The Zululand shells are remarkable for the great relative height 
attained by several individuals. In one of the specimens before 
me (T.M. 1894) the height exceeds the length by a considerable 
amount (1: 1-48); taken by itself one would be inclined to regard the 
specimen as distinct from P. sphaeroidea, but in the remaining 18 
specimens, however, the ratio of length : height works out at from 
1: 1-11 to 1: 1-21, which is well within the range given above for 
P. sphaeroidea. In proportions there is a very striking resemblance 
of some of the shells before me to the shell from the Upper Greensand 
of Haldon (Upper Albian) figured as Protocardia sp. by Woods (1908, 
p. 196, pl. xxxi, fig. 4), and it is unfortunate that the ornamentation 
of the latter is so poorly preserved; this shell is said by Woods to 
resemble in shape the higher forms of P. sphaeroidea (from the Lower 
Greensand), but to have a narrower and more curved umbo and a more 
distinct carina. The record made by Jukes-Browne (Woods, loc. cit.) 
of P. sphaeroidea, from the zone of Pecten asper (L. Cenomanian) of 
Wiltshire, was possibly based on a shell of this type. On the whole, 
the umbonal region is narrower and more pointed in the Zululand 
shells than in the two Lower Greensand shells figured by Woods, 
and there is a closer approximation to the Upper Greensand shell in 
the form of the umbonal region, but there is an approach to the 
broader umbo of P. sphaeroidea in some specimens. In the type 
specimen of P. rothpletzi the umbo is broader than in any of the 
Zululand shells. 

The concentric ribbing of the English and Zululand shells is essen- 
tially the same in character, flat, broad ribs with narrow grooves 
separating them. In the examples of P. sphaeroidea, figured by 
Woods, there are 5-7 ribs in 10 mm. in the middle part of the shell, 
judging from the figures which are presumably approximately 
natural size. In the type of P. rothpletzi, again judging from the 
figure, there are about 7-8 ribs in 10 mm., the grooves being slightly 
wider than in Woods’s figures. In the Zululand shells the number 
of ribs is usually about 8 or 9 in a distance of 10 mm. at about 50-60 
mm. from the umbo, but in one instance (S.A.M. 10788) the number 
of ribs in that distance is only 5. In P. sphaeroidea and in the 
Zululand shell 8.A.M. 10788 the ribs are remarkably broad and flat, 
while in the majority of the Zululand shells the ribs are relatively 


Lower Cretaceous Lamellibranchia from Northern Zululand. 381 


narrower and separated by wider grooves, and a little more rounded, 
thus resembling those of P. rothpletz. 

It would appear from the above facts that there are no very striking 
features which can be used as a means of discriminating between the 
English, Hast African, and Zululand shells, and it may well be the 
case that we are dealing here with sets belonging to a single, if 
somewhat variable, species. The peculiarly short Zululand shell 
(T.M. 1894) mentioned above can be safely regarded as an extreme 
case, an abnormal individual in which great relative height and 
more pointed umbonal region are correlated, since no less than 18 
associated specimens of the same general character, and undoubtedly 
conspecific with it, have relatively longer shells. It should be pointed 
out that the characters of P. sphaeroidea were judged on the basis of 
the two figures given by Woods, and that an examination of an 
adequate set of this species might disclose grounds for a more em- 
phatic expression of opinion as to relationship with the Zululand 
shells. 

P. sphaeroidea and P. rothpletzi are Lower Aptian and Neocomian 
respectively, while the Zululand shells * occur in association with 
Albian species. The imperfectly preserved shell figured by Woods 
from Haldon is Upper Albian. 

A feature of the Zululand shells f not mentioned by Woods for 
P. sphaeroidea is the presence of a tooth-like tubercle within the left 
valve at the postero-ventral corner; with the growth of the valve the 
tubercle gives rise to a very slightly or scarcely raised internal rib 
corresponding in position to the posterior carina. The tubercle fits 
into a slight embayment of the corresponding margin of the right valve, 
the embayment being rounded, about 1 to 1-5 mm. in depth, and 
distinctly visible when the valve is viewed from beneath. 

Dietrich (loc. cit.) included P. rothpletzi and P. sphaeroidea in 
a new sub-genus Tendagurium, under Cardium, the type of the 
sub-genus being another Hast African species C. propebanmianum 
Dietrich, which is ornamented only with fine growth lines over 
the whole surface. It is doubtful whether these two species are 
closely related to the latter, and they are accordingly retained under 
Protocardia. 


* Etheridge (1907, p. 77, pl. iii, figs. 6, 7) gave the new name Protocardiwm ? 
cinctutum to a very imperfect shell from the Umsinene River and suggested 
affinity with P. sphaeroidea, probably rightly. 

+ Denudation of the carina gives rise to a double impressed line in many of 
the specimens studied. 

VOL. XXXI, PART 3. 31 


‘ 


382 Annals of the South African Museum. 


Dimensions of representative Zululand shells, expressed in 


millimetres: ' 
T.M. T.M. T.M. S.A.M. S.A.M. 
1790. 1796. 1894. 10788. 10789. 


Length 5/5) 52 5b ae eae 15 
Height : ws 60 Gl 7 S260 81 84 
Thickness of one 

valve : 4 e238 20 21 28 29 33 


Genus CarpDium Linnaeus 1758. 
Cardium rogersi sp. nov. 
(Plate XLIX, figs. 3-8.) 


Material.—The species is founded on about 28 specimens, the 
majority more or less complete. For the most part the specimens 
are well covered with matrix, but a number were cleaned up sufficiently 
well to show the external features and the hinge. Mr. H. F. From- 
murze collected several specimens on the Mfongosi stream (S.A. Mus. 
10791-10796). Dr. 8S. H. Haughton and Dr. A. W. Rogers collected 
the species as follows: Mfongosi stream, Z2 (17), Z3 (1); Myesa 
stream, M1 (2). The holotype (S.A.M. 10791) and figured speci- 
mens are for the most part taken from the earlier collection of 
Mr. Frommurze, in that the general preservation is superior, and in 
spite of the fact that the horizon is not precisely recorded. 

Description.—Shell moderate in size, about as high as long, relatively 
stout, very convex, slightly inequilateral. Umbones very prominent, 
highly incurved. Hinge line long, nearly as long as the greatest 
length of the shell, nearly straight. Anterior margin convex, passing 
obliquely downwards in a broad curve to merge gradually into the 
convex ventral margin. Posterior somewhat truncate, the posterior 
margin nearly vertical, the upper and lower parts convex, the middle 
part concave, meeting the ventral margin in a broadly rounded 
postero-ventral angle. Valve evenly convex, the posterior slope 
becoming steeper than the anterior, with the postero-dorsal corner 
forming a wing which is separated from the posterior slope by a broad 
concavity. 

Dentition relatively strong, the two cardinals in the left valve close 
together under the umbo, the lower or anterior cardinal conical and 
prominent, the upper or posterior cardinal smaller; laterals in left 
valve prominent, oval in section, the anterior more prominent than 
the posterior ?, situated at some distance from the cardinals. Anterior 
and ventral margins crenulate within, with numerous short grooves 


Lower Cretaceous Lamellibranchia from Northern Zululand. 383 


transverse to the margins; at the postero-ventral angle these pass 
into stronger marginal incisions, the posterior margin being coarsely 
denticulate. 

Ornamentation essentially radial, that of the posterior slope being 
considerably coarser and different from that of the rest of the valve, 
the two areas being sharply demarcated after the period of early 
middle growth, the line of demarcation being a little more than one- 
third of the length of the valve from the posterior margin. Flank 
with numerous incised radiating lines, separated by relatively wide 
flat interspaces; on partially decorticated surfaces a very fine con- 
centric ribbing comes in, giving a minutely reticulate surface. Towards 
the anterior the radial incisions widen and the shell becomes orna- 
mented with very faint radial riblets. The posterior area is orna- 
mented with about 20 strong, raised, sharp, radial ribs, separated by 
deep U-shaped grooves, the ribs corresponding to the teeth on the 
posterior margin. 


Dimensions, in millimetres: 
S.A.M. S.A.M. S.A.M. From 
10791. 10792. 10793. Z2. 


Length Sot 31 30 38 
Height ; 7) 39 DOs eae ears 39 
Length of hinge (approx.) . 27 25 26 27 
Thickness of one valve L eonalh 12 16 al 


Remarks.—The species is quite distinct from any South African 
Cardiidae hitherto described. 


Famity VENERIDAH. 
Genus PrycHomya Agassiz 1842. 
Ptychomya robinaldina (d’Orbigny). 
(Plate XLIX, figs. 1, 2.) 


1844. Crassatella robinaldina d’Orbigny: Pal. Frang. Terr. Crét., 
volta, 7: (5, pl. celxiv, figs. 10-13. 

1907. Ptychomya robinaldina (d’Orb.): Woods, Cret. Lam. Engl., 
Ol Th pid, pl xxvir, hes. 25-26. 

The record of this species from Northern Zululand is based on a 
single specimen (S. Afr. Mus. 10812) collected by Mr. H. F. From- 
murze on the Mfongosi stream, the precise horizon not being noted. 
The specimen consists of the two valves firmly closed. The whole 
surface is denuded, the dorsal part badly so, so that the umbonal part 


384 Annals of the South African Museum. 


is deceptively inconspicuous and some allowance must be made for 
rounding of the marginal angles. Nevertheless the salient character- 
istics of form and sculpture are unmistakably indicated, and I see no 
reason for regarding this shell as distinct from the Lower Greensand 
specimens described and figured by Woods. The shell is of the same 
size and general proportions as in the English examples and is orna- 
mented in the same way. The posterior part of the shell is very much 
worn, but the ribs in that part were clearly stronger and more widely 
separated than elsewhere, while the postero-dorsal margin bears 
unmistakable traces of coarse ridges and crenulation. The main 
portion of the flank bears ribs of exactly the same character and about 
the same in number as the English shells. Towards the anterior 
a chevron arrangement of the ribs is just discernible in spite of 
wear. 

There are certain features which might suggest some distinction 
between this shell and the two specimens figured by Woods, but these 
appear to be of little importance. The umbones appear to be less 
conspicuous and there is a greater rounding of the postero-dorsal 
angle, while the valves appear to be less convex when seen from above; 
to a large extent these differences may be assigned to the denuded 
character of the specimen. The anterior part is very slightly narrower 
and more produced than in the English shells, the umbones being 
placed a little farther back. The chevron arrangement of the anterior 
ribs forms a pattern which does not precisely agree with that of either 
of the shells figured by Woods, but the latter has pointed out that 
there is a considerable range of variation in this respect, as in other 
species of lamellibranchs having an analogous sculpture. 

P. complicata (Tate) (1867, p. 160, pl. ix, fig. 8), referred to the 
genus by Dames (Kitchin, 1908, p. 27), was founded on a very small 
shell from the Sundays River Beds which appears to differ in the 
details of sculpture. 

P. kitchen Lange (1914, p. 242, pl. xvu, fig. 6), from the Trigonia 
schwarzi-Bed in Tanganyika, is less elongate, and finely ribbed in 
the posterior part, besides differing in other details. Dietrich (1933, 
p. 53) has shown that this species is identical with the previously 
described P. hawchecorner Miller from the same area, and has placed 
it as a variety of P. robunaldina. 

According to Dietrich (loc. cit.) the genus ranges from the Neocomian 
to the Upper Cretaceous, and Kitchin (1926, p. 460) has pointed out 
that it forms a characteristic element of the southern Neocomian 
faunas. 


Lower Cretaceous Lamellibranchia from Northern Zululand. 385 


FAMILY uncertain. 
Genus THETIRONIA Stoliczka 1871. 
Thetiroma sp. 


Imperfect shells apparently belonging to the same species were 
collected from L12 (1) on the Lombagwenya stream, and Z24 (1) 
south of Mokatini. 


Famity SAXICAVIDAE. 
Genus Panore Ménard de la Groye 1807. 


(= Panopea and Panopaga of authors). 
Panope gurgitis (Brongniart). 


(PlatemipeinesssIeto.16,.8, 9.) 
(Figs. 2-4, 7, 10, 11 from the Uitenhage Series.) 


1850. Anoplomya lutrarva Krauss: Unt. Kreide Kaplandes, p. 447, 
Toll, sxlbeamly wea Ike 

1907. Myopsis? africana Htheridge: 3rd Rep. Geol. Surv. Natal, 
TO. Gli, Ole ty, seas), ILO) 

1907. Glycimeris? griesbacha Etheridge: Ibid., p. 81, pl. iii, 
fig. 9 (non Glycymeris griesbachi Newton, Arcidae). 

1908. Pleuromya lutraria (Krauss): Kitchin, Ann. 8S. Afr. Mus., 
Ol, Wilh JOO. De, MS 

1909. Pleuromya africana (Etheridge): Newton, Trans. Roy. Soc. 
S. Afr., vol. i, p. 84, pl. vii, figs. 1-3. 

1909. Panopea gurgitis (Brongniart): Woods, Cret. Lam. England, 
vol. 11, p. 222, figs. on pls. xxxv, XXXVl. 

Non Pleuromya africana (Etheridge): Rennie, Ann. 8. Afr. Mus., 
vol. xxvii, p. 186, pl. xx, figs. 8-10, 1930. 


Material.—The collections include about 30 specimens indistinguish- 
able from this well-known European species. These were collected by 
Dr. 8. H. Haughton and Dr. A. W. Rogers at the following localities: 
Myesa stream, M1 (several, imbedded in hard limestone); Lombag- 
wenya stream, Ll (1), L3 (2), L7 (4), L9 (2), Lil (1); Mfongosi 
stream, Z6 (1), Z8 (1), Z9 (4), Z11 (1), Z12 (3), Z14 (1); Pongola 
River below junction with Mfongosi, Z19 (1), Z20 (1). 

Remarks.—The shells before me from the Pongola area do not 
appear to be distinguishable on external characters from certain 
forms which Krauss, Etheridge, and Newton have described and 


386 Annals of the South African Museum. 


figured under other names from South African Lower Cretaceous 
deposits. I have been unable to find satisfactory reasons for separat- 
ing the species named by these authors from the well-known European 
Panope gurgitis, accepting this species in the wide sense adopted by 
Woods in his monograph of the Cretaceous Lamellibranchia of 
England. 

Krauss (loc. cit.) long ago described and figured certain shells from 
the Zwartkops River, Sundays River Beds, as Anoplomya lutraria 
gen. et sp. nov., said to have the habit of a Lutraria (Mactridae), 
but to be close to Panope, though differing from the latter in certain 
respects. Krauss was led to institute a new genus for the species 
on the grounds that the cardinal teeth were assumed to be absent; 
a peculiar strong crenulation of the dorsal margin of each valve near 
the umbo was described and figured. I have not been able to find 
specimens of this species, which are not uncommon in collections, 
showing the hinge completely exposed, but from a comparison of 
Krauss’s figures and fine specimens of Panope natalensis Woodw. 
from a raised beach at Port Elizabeth, I have no doubt that Krauss’s 
specimens possessed the hinge structure characteristic of the genus 
Panope. In that genus the solitary prominent cardinal tooth of each 
valve is situated close to the hinge margin and immediately below 
the umbo, while posterior to it the strong elongate nymphal structure 
projects well beyond the hinge margin; the crenulation of the dorsal 
margin figured by Krauss for each valve corresponds exactly in out- 
line with the projecting tooth and nymph of Panope, and Krauss 
was clearly in error in his interpretation of these structures; in his 
fig. 1 b, the right valve is apparently slightly out of place, so that the 
right tooth and nymph appear to overlap the margin of the left 
valve. According to Kitchin (1908, p. 159) and Htheridge (1907, 
p- 82) Anoplomya has been quoted as a synonym of Pleuromya, but 
Kitchin, Newton (1909, p. 85), and other authors have included 
Panope gurgitis in Pleuromya. According to Woods (1909, p. 225) 
Panope is distinguished from Pleuromya by having a well-developed 
tooth, contiguous but not overlapping dorsal margins, and a well- 
developed external ligament; in a specimen of A. lutraria before me 
(A.M. 3840) the hinge of the right valve is sufficiently exposed 
to show a deeply impressed ligament groove and a projecting but 
relatively short nymph, which, with the interpretation of Krauss’s 
figures given above, supports reference of Anoplomya to Panope. 
It is significant that Kitchin (loc. cit., p. 158) associated A. lutraria 
with forms referred by Woods to P. gurgitis, and I have failed to 


a 


Lower Cretaceous Lamellibranchia from Northern Zululand. 387 


find any distinction between a number of specimens from the Sundays 
River Beds (in Albany Museum Coll.) and the numerous examples 
figured by Woods from the lower Greensand of Atherfield. Both 
the typical and plicate (var. plicata (J. de C. Sow.)) types are re- 
presented, the distinction being insufficient to warrant recognition 
of two species, since the forms grade into one another, as Woods 
has pointed out. 

Glycumeris? griesbachi Etheridge (1907, p. 81, pl. iii, fig. 9), from 
the Umsinene River, not to be confused with another Zululand shell 
of the same name in Glycymeris da Costa (G. griesbachi Newton, 
1909, p. 36), was founded on very imperfect material that could very 
well have been referred to P. gurgitis. 

Myopsis? africana Ktheridge (1907, p. 81, pl. i, figs. 9, 10), from 
the same locality, was distinguished as a separate species from the 
above in part by reason of the shorter anterior and more distinct 
anterior carina. The figured specimen is very imperfect, and it 
would appear that the postero-dorsal margin is broken away, giving the 
specimen a spurious appearance of narrowing towards the posterior, 
an appearance which in part led the author to identify with it two 
shells from the Upper Senonian of Pondoland (Rennie, 1930, p. 186, 
pl. xx, figs. 8-10). The latter species is certainly not a form of 
Panope gurgitis, the posterior being narrow and almost pointed, and 
reference to Pleuromya appears to have been correct. The type of 
M.? africana is ornamented with radial lines of granules as in some 
of the shells of P. gurgitis figured by Woods, and reference to the 
latter is highly probable. The shell figured by Newton (1909, p. 84, 
pl. vii, figs. 1-3) from the Manuan Creek as Pleuromya africana 
(Hth.) was compared with forms included in P. gurgitis by Woods, 
and appears to be indistinguishable from the latter species. 

It is apparent from Woods’s monograph that Panope gurgitis is 
a very variable species and that a number of unnecessary names 
have been proposed for European material from different localities. 
The variation is in respect of the strength of the concentric folds, 
and in the details of outlines and form. Fine radial granulation, 
formerly used to distinguish P. neocomiensis from P. gurgitis, appears 
in good specimens of the latter. The South African species referred 
to above cannot be distinguished in form from this variable species, 
and fine radial lines of granules were noted by Etheridge and Newton 
in some of the shells before them. In the present collection the 
material ig either weathered or embedded in hard limestone, but 
the reference to P. gurgitis is justified on external characters; the 


388 Annals of the South African Museum. 


specimens are fairly typical, those from L7, L9, and Lill being 
somewhat plicate and relatively short. 

In Europe the species ranges from Neocomian to Upper Albian. 
P. lutraria from the Sundays River Beds is Upper Valanginian. In 
the collection of the Transvaal Museum there are two fine specimens 
of P. gurgitis (T.M. 1261, 111) from Catuane, the horizon being 
Upper Albian (Spath, 1925, p. 180), one specimen being very typical, 
the other more distinctly plicate. 


GASTROPODA. 


The comparative scarcity of Gastropods is a feature of the faunas. 
Small shells belonging to Dicroloma are found at several horizons, 
and those from localities L9, Z1, and Z2 appear to belong to the 
same species. The well-characterised species Turritella manuanensis 
Newton (1909, p. 25, pl. vii, figs. 16, 17), described from the Manuan 
Creek, is represented in the collections by a number of specimens 
from the neighbourhood of the Mfongosi stream: Z16 (many, more 
or less weathered), Z19 (10), Z20 (2); the species is said by du Toit 
(1926, p. 318) to be from the Albian. Rather imperfectly preserved 
shells, probably identical with the form described by Newton (zbid., 
p- 30, pl. vii, figs. 6-8) from the Manuan Creek as Avellana cf. 
encrassata (J. Sow.), were found at two localities, L1 (1) and Z19 (2), 
at the latter locality associated with Albian species. On the Mfongosi 
stream a number of specimens similar to Pterodonta inflata d’Orb. 
were collected at Z2 (12), but whether this Cenomanian species is 
really represented is doubtful. 


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Trigoniidae,” Pal. Nav. Nas. Mus. Bloemfontein, deel 1, stuk 1, pp. 1-38, 
pls. i-vii. 

vAN Hoepen, E. C. N. (1929 a).—Internat. Geol. Congress, Session XV, guide book, 
Durban-Zululand excursion, pp. 10-18. 


Lower Cretaceous Lamellibranchia from Northern Zululand. 391 


van Hoxpsn, HE. C. N. (1931).—* Voorlopige beskrywing van enige Soeloelandse 
Ammoniete,” Pal. Nav. Nas. Mus. Bloemfontein, deel 1, stuk 2. 

Wank, B. (1926).—“‘The Fauna of the Ripley Formation, Tennessee,” U.S.G.S. 
Prof. Paper, No. 137. 

Weaver, C. E. (1931).—“‘ Palaeontology of the Jurassic and Cretaceous of West 
Central Argentina,’ Mem. Univ. Washington, No. | (not seen). 

Woops, H. (1899-1913).—Mon. Cretaceous Lamellibranchia England, Mon. Pal. 
Soc. 

Woops, H. (1906).—“The Cretaceous Fauna of Pondoland,” Ann. S. Afr. Mus., 
vol. iv, pt. vii, pp. 275-350. 

ZitTEL, K. (1913).—Text-book of Palaeontology, English edition, ed. C. R. 
Eastman. 


EXPLANATION OF PLATES. 


The figures are approximately natural size unless otherwise stated; in the 
majority of cases the figures are very slightly reduced. 


PuaTE XXXVII. 
FIGS. PAGE 
1-3. Cucullaea (Megacucullaea) spp. : : : : ‘ : . 3808 


Lombagwenya stream, Northern Zululand. 


1. Left valve from L12. 
2. Fragment of anterior part of large left valve from L9. 
3. Weathered and crushed left valve from L12. 


Plate XX XVII. 


Ann. S. Afr. Mus., Vol. XX XT. 


eill & Co., Ltd. 


NV 


HenOtOore ds). Vi. ly, R. 


LAMELLIBRANCHIA FROM NORTHERN ZULULAND. 


Cucullaea (Megacucullaea) spp. 


yy 
bal 
a - 
= % 
2 
- 
a P . 

w- 7 
: ’ 

= ' » 

4 “es 
= 
‘ 
& 
: 
‘ 
~ 
a x 
=) 
~ 
vo oD a 


OAs Nie Se AIR OTE oe he Sy a ees 


FIGS. 
1-3. Trigonia (Megatrigonia) obesa (van Hoepen) 
Mfongosi stream, Northern Zululand. — 
1. Specimen from Z3, dorsal view. 
2. The same, anterior view. 
3. ,,. right valve in lateral view. 


, 4 a ht oe 


‘ a 
i. ee = 


ee Te ee eh tle Ay ok 


Ann. S. Afr. Mus., Vol. XX XI. Plate XX XVIII. 


Eno: J.V.L. R. Neill & Co., Ltd. 


LAMELLIBRANCHIA FROM NORTHERN ZULULAND. 


Trigonia obesa. 


PuaTE XXXIX. 
FIGS. PAGE 
1, 2. Trigonia (Megatrigonia) obesa (van Hoepen) . : : ; . one 
Mfongosi stream, Northern Zululand. 
1. Left valve from Z3 seen in lateral view. 
2. Right valve from Z3 seen in lateral view. 


Ann. S. Afr. Mus., Vol. XX XI. 
Plate XOe RIX. 


Eigigend. V.L. R. Neill & Co., Ltd. 
LAMELLIBRANCHIA FROM NORTHERN ZULULAND. 
Trigonia obesa. 


32 


Puate XL. 
FIGS. 


1-3. Trigonia (Megatrigonia) conocardiiformis (Krauss) 


Sundays River Beds, Uitenhage Series, Cape E 


1. Specimen from Sundays River, collec 
Atherstone, Albany Museum 738, 1 
lateral view. : 

2. The same, dorsal view. 

a: anterior view. 


99 


o 
-_ 
3 


ArnmerseeAtr, Mus., Vol. XX XI. 


Plate: Xi. 


Co., Ltd. 


Neill & 


SERIES. 


wy) 


ITENHAGI 


+ 
} 


l 
for 


4) 
1 


FROM THE 


LLIBRANCHIA 


wy) 


LAMI 


LLB. Te eee hee 


Photo 


"MAUS, 


card 


rigonia conoc 


4h 


Pirate XLI. 
FIGS. PAGE 


1-4. Trigonia (Lotrigonia) haughtoni sp. nov. . 2 : : ‘ . 340 
Myesa stream, Northern Zululand. 


1. Hand specimen, with holotype and paratypes from M1; 
the holotype, the larger right valve on the right side of 
the specimen; paratypes, a left valve below and a 
damaged right valve on the left side of the specimen; 
natural size. (Figs. 2-4 are enlargements of these.) 

2. Paratype, right valve, dorsal view to show area and 
escutcheon, x 2:65. 

3. Paratype, left valve, slightly oblique dorsal view to show 
area and escutcheon, x 2:5. 

4. Holotype, right valve, seen in lateral view, x 2. 


5, 6. Trigonia (Rutitrigonia) pongolensis sp. nov. : : I : - 309 
Mfongosi stream, Northern Zululand. 


5. Paratype from Z2, left valve seen in lateral view. 
6. The same, dorsal view. 


Ann. S. Afr. Mus., Vol. X LI. Plate XLI. 


eno: J. V.L. R. Neill & Co., Lid. 


LAMELLIBRANCHIA FROM NORTHERN ZULULAND. 
1-4. Trigonia haughtoni. 5, 6. Trigonia pongolensis. 


Prats XLII. 
FIGS. PAGE 
1-4. Trigonia (Pleurotrigonia) blanckenhorni R. B. Newton ‘ : . 363 


South of Mokatini, Northern Zululand. 


1. Left valve from Z25, umbonal region enlarged x 2 approxi- 
mately, to show beaded marginal carina and traces of 
costellae on the area. 

2. The same, anterior view, natural size. 


oF a seen in lateral view, natural size. 
4. 3 oblique dorsal view, x 2 approximately. 
5-7. Trigonia (Rutitrigonia) pongolensis sp. nov. : : ‘ : . 309 


Mfongosi stream, Northern Zululand. 


5. Holotype, left valve from Z2, seen in lateral view. 
6. The same, dorsal view. 


7 


ie a hinge, x 2. 


Ann. S. Afr. Mus., Vol. XX XI. Plate XLII. 


Photo: J. V. 1. R. Neill & Co., Lid. 
LAMELLIBRANCHIA FROM NORTHERN ZULULAND. 


1-4. Trigonia blanckenhorni. — - 5-7. Trigonia pongolensis. 


— a See —— —- —— —— ——— ee — a ee = 
— oe RR ———— Nal ae ee = artery a ——= = = a = = 
—- - en ee ee ea = : ——— i 


oe 
we mi 


4 
va 
‘ 
aT 
Ad 


Prate XLITI. 
FIGS. PAGE 
1-3. Trigonia (Steinmanella) hennigi Lange : : , ; . d47 


Mfongosi stream, Northern Zululand. 


1. Specimen S. Afr. Mus. 10786, left valve seen in lateral 
view. 

2. The same, dorsal view. 

oF ps anterior view. 


4. Pholadomya luynesi Lartet ; : . : : ; <) paned. 
Pongola River, Northern Zululand. 


4. Specimen from Z19, left valve seen in lateral view, natural 
SIZe. 


Ann. S. Afr. Mus., Viole xXOxOx: Plate XLIII 


Photo: J. V. L. R. Neill & Co., Ltd, 


LAMELLIBRANCHIA FROM NORTHERN ZULULAND. 
1-3, Trigonia hennigi. . 4. Pholadomya luynest. 


PuatE XLIV. 
FIG. PAGE 
1. Trigonia (Steinmanella) hennigi Lange ‘ ‘ : ; . . 347 


Lombagwenya stream, Northern Zululand. 


Specimen from L14, right valve seen in lateral view. 


2. Cucullaea (Megacucullaea) kraussi Tate : : : : . 3805 
Sundays River Beds, Uitenhage Series, Cape Province. 


Portion of right valve from Zwartkops River, Albany 
Museum 792, collected by Dr. W. G. Atherstone, to show 
radial ornamentation. 


Ann. 8. Afr. Mus., Vol. XX XI. Plate XLIV. 


Photo: J. V. L. R. Neill & Co., Ltd. 


LAMELLIBRANCHIA FROM NORTHERN ZULULAND AND THE UITENHAGE SERIES. 
1. Trigonia hennigi. 2. Cucullaea (Megacucullaea) kraussi. 


iat) ah 
we oF 
‘4 ; 
} 

y 


PLATE XLV. 
FIGS. PAGE 
1-3. Trigonia (Sphenotrigonia) frommurzet sub-gen. et sp. nov. . ; . 367 


Mfongosi stream, Northern Zululand. 


1. Holotype, 8. Afr. Mus. 10586, left valve seen in lateral 
view. 

2. The same, dorsal view. 

3. The same, anterior view (the gentle lunular depression 
does not show up in this photograph). 


Plate XLV. 


Ann. 8S. Afr. Mus., Vol. X X XI. 


PIT “OD YP [AN 


CNWTINTOZ 


“VAZINUULOLf DIUObLA T, 


NUYHHLYON 


WOU VIHONVUYPITIONVI 


“UT A LF 0N0Ug 


ae 


a 


, 
/ 
ry 
, 
2 


cae 


vee PTs ite fies 
Ve Le ee Tet 


ad a i L 


vee 


Pirate XLVI. 
FIGS. PAGE 
1-4. Trigonia (Sphenotrigonia) frommurzei subgen. et sp. nov. . : . 367 


Mfongosi stream, Northern Zululand. 


1. Left valve, S. Afr. Mus. 10591, seen in lateral view; a 
considerable part of the posterior end is missing. 

. Anterior part of an immature left valve from Z7, interior 
view to show the dentition. 

. Specimen S. Afr. Mus. 10588, anterior view. 

4. Same specimen as fig. 3, dorsal view; a considerable part 


bo 


ivy) 


of the posterior end is missing. 


Ann. S. Afr. Mus., Vol. XX XI. Plate XLVI. 


Photo: J.V L.R. Neill & Co., Ltd. 
LAMELLIBRANCHIA FROM NORTHERN ZULULAND. 


: Trigoma frommurzet. 


— a ee oe = —— 
— 2 A a ASA Aap 


Prats XLVI. 
FIGS. 
1-4. Trigonia (Sphenotrigonia) frommurzei subgen. et sp. nov. 


Mfongosi stream, Northern Zululand. 


1. Anterior part of a mature left valve from Z13, interior 
view to show the dentition; a considerable part of the 
posterior end is missing. 

2. Anterior part of an immature left valve from Z7, the same 
specimen as Plate 10, fig. 2, oblique dorsal view to 
show the juvenile costae and beaded carinae, x 2 
approximately. 

3. Right valve from Z8, anterior view. 

4. The same specimen as fig. 3, interior view. 


PAGE 


367 


Ms. Afr. Mus., Vol. XXXI. 
Plate XLVII. 


Photo: J.V.L.R. Neill & Co., Ltd. 
LAMELLIBRANCHIA FROM NORTHERN ZULULAND. 


Trigonia frommurzer. 


— Jf : 
ssa ee 


4 


] s 
¥ ‘ i 
%, % i f 
i a es ny 
~ ag yt : ’ fl if 
iy : i j : 
at on 7 ie 
, 
& 
; ‘ 
4s 
5 * 
. ‘ 
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‘ 
OL. Xx, PART 3. 
% 
i i 
a, : = 
feat | 
i} 
i I 
dan 


© 


33 


Puate XLVIII. 
PIGS. PAGE 
1-6. Protocardia ct. sphaeroidea (Forbes) . : : : : : - 319 


Mfongosi stream, Northern Zululand. 


1. Left valve from Z19, seen in lateral view. 

2. Same specimen as fig. 1, interior view; note tubercle at 
margin, postero-ventral corner. 

3. Same specimen as fig. 1, dorsal view. 

4, Right valve, Tvl. Mus. 1796, seen in lateral view. 

Be »  Ivl. Mus. 1894, seen in lateral view. 

A » trom Z19, seen in lateral view. 


oo 


Ann. S. Afr. Mus., Vol. XX XI. 
Plate XLVITI. 


Photo: J. V. L. R. Neill & Co., Ltd 


LAMELLIBRANCHIA FROM NORTHERN ZULULAND. 
Protocardia cf. sphaeroidea. 


7," 


FIGS. 


Puate XLIX. 


1, 2. Ptychomya robinaldina (d’Orbigny) . 


Mfongosi stream, Northern Zululand. 


Ihe 


Specimen S. Afr. Mus. 10812, left valve seen in lateral 
view; very worn specimen, the chevron arrangement of 
the anterior ribs scarcely but unmistakably preserved. 


2. The same, dorsal view. 


3-8. Cardium rogersi sp. nov. 


Mfongosi stream, Northern Zululand. 


3. 


4. 
5. 


=~] 


Holotype, a left valve, S. Afr. Mus. 10791, seen in lateral 
view. 
Left valve from Z2, interior view. 


Left valve, S. Afr. Mus. 10792, interior view; the sub- 


umbonal portion of the hinge has been fractured and 
depressed slightly. 


. Same specimen as fig. 5, seen in lateral view. 
. Holotype, same specimen as fig. 3, dorsal view. 
. Right valve, S. Afr. Mus. 10793, seen in lateral view; a 


considerable part of the ventral and posterior borders 
missing. 


9. Protocardia cf. sphaeroidea (Forbes) 


Mfongosi stream, Northern Zululand. 


9. Large left valve, S. Afr. Mus. 10788, seen in lateral view. 


PAGE 
383 


382 


379 


Ann S. Afr. Mus., Vol. XX XI. 
Plate sxUL exe 


MOLOYS Jia Vie Tre Be Neill & Co., Lid. 


LAMELLIBRANCHIA FROM NORTHERN ZULULAND. 


1, 2. Ptychomya robinaldina. 3-8. Cardium rogersi. 
9. Protocardia cf. sphaeroidea. 


s 
;% 
& 
’ 
7 
: 
bp 
uF 
‘ 
i 
° 
Y 
, 
. 
sf 
Ya 


FIGS. 


PuatTE L. 


1-11. Panope gurgitis (Brongniart) . 


Figs. 2-4, 7, 10, 11 are specimens of “‘ Anoplomya lutraria”’ Krauss 
from the Uitenhage Series. 


Wigs: 1.25556; 15..0 from Northern Zululand. 


is 


ie) 


a | 


ie 


Specimen from the Lombagwenya stream, Northern 
Zululand, L7, right valve seen in lateral view. 


. A. lutraria, specimen from Zwartkops River, Uitenhage 


Series, collected by Dr. W. G. Atherstone, right valve 
seen in lateral view; probably a topotype of Krauss’s 
species. In Alb. Mus. Coll. 


. The same specimen as fig. 2, dorsal view. 
. A. lutraria, specimen from Redhouse, Zwartkops River, 


Uitenhage Series, collected by Mrs. T. V. Paterson, left 
valve seen in lateral view; probably a topotype of 
Krauss’s species. In Alb. Mus. Coll. 


. Specimen from the Lombagwenya stream, Northern 
Zululand, £7, portion of right valve seen in lateral view. 
. Right valve from Mfongosi stream, Northern Zululand, 


Z9, seen in lateral view. 


. A. lutraria, specimen from Zwartkops River, Uitenhage 


Series, collected by Dr. W. G. Atherstone, right valve 
seen in lateral view; probably a topotype of Krauss’s 
species. In Alb. Mus. Coll. 


. Specimen from the Mfongosi stream, Northern Zululand, 


Z12, left valve seen in lateral view. 


. The same specimen as fig. 8, dorsal view. 
10. 


A. lutraria, specimen from Uitenhage Series, probably 
part of set Albany Museum 3634-7 collected by Prof. 
E. H. L. Schwarz at Addo on the Sundays River, 
seen in lateral view. 

The same specimen as fig. 10, dorsal view. 


PAGE 
385 


Ann. S. Afr. Mus., Vol. XX XT. 


Plate L. 


Photo: J. V. L. R. Neill & Co., Lid. 


LAMELLIBRANCHIA FROM NORTHERN ZULULAND AND THE UITENHAGE SERIES. 
Panope gurgitis. 


oe 


Puate LI. 


. 


_ Mfongosi stream, Northern Zululand. Se 
1. Left valve, interior view, S. Afr. Mus. 10807. 
2. The same, seen in lateral view. ; 


1, 2. Gervillia dentata Krauss 


FIGS. 


| 


ty 


Ann. S. Afr. Mus., Vol. XX XI. 


JOKD & ti. Wee Ibe, IF. 


LAMELLIBRANCHIA FROM NORTHERN ZULULAND. 
Gervillia denatata. 


Neili & Co., Ltd. 


Puate LII. 
FIGS. PAGE 
1-3. Gervillia dentata Krauss . : : ; : : ; : . “Si0 


Mfongosi stream, Northern Zululand. 


1. Specimen S. Afr. Mus. 10802, right valve seen in lateral 
view. 

2. Specimen S. Afr. Mus. 10804, right valve seen in lateral 
view. 

3. The same specimen as fig. 2, dorsal view. 


Ann. s. Afr. Mus., Vol. XX XI. 


Photo: J. V. L. R. 
LAMELLIBRANCHIA FROM NORTHERN ZULULAND. 


Gervillia dentata. 


Piste uu: 


Neill &: Co., Ltd. 


Ve fag aes 


_ 
a - : é 7 74 e 
ws 7 ha a aan 5 y . 
} r 1 at ‘ view ore ea - 
Xe i Piel abe. 
; i i , ao al 
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Puate LITI. 
FIGS. PAGE 
1-3. Gervillia dentata Krauss . ; ; : ‘ : : : 5 Bi 


Sundays River Beds, Uitenhage Series, Cape Province. 


1. Specimen from the Sundays River, collected by Dr. W. G. 
Atherstone, Albany Museum 822, left valve seen in 
lateral view, reduced x 0-83 approximately, showing 
the demarcated anterior area. 

2. The same, dorsal view, reduced x 0-83 approximately. 

3. Specimen from the Sundays River, collected by Dr. W. G. 
Atherstone, Albany Museum 810, anterior portion of 
left valve, showing the demarcated anterior area. 


Ann. S. Afr. Mus., Vol. XX XI. 
Plate LITT. 


Photo: J. V. L. R. Neill & Co., Ltd. 


LAMELLIBRANCHIA FROM THE UITENHAGE SERIES. 


Gervillia dentata. 


-? 

a) 
f i 
i 

i} 

. 

= 
| a 
i + 
| 

F 
\* 
| 
| 
| 
I 
-- 


: PLatEe LIV. 
FIGS. 
1, 2. Gervillia dentata Krauss : $ F 4 
Sundays River Beds, Uitenhage Series, Cape 
iz 1. Right valve from the Sundays River, col 
| 7 Atherstone, Albany Museum 823, seen 
| the posterior part missing. . ; 
2. The same, interior view. 


i} 
‘ 
/ 
| 
t 
| 
| 
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} 
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c 
= 
ee 
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C: r 
7 fag Ate 
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Geeta nr” 


| . 8. Afr. Mus., f : 
Ann. S. Afr. Mus., Vol. XX XT Plate LIV. 


whoto: J. V. L. R. Neill & Co., Lid. 


LAMELLIBRANCHIA FROM THE UITENHAGE SERIES. 
Gervillia dentata. 


PuatEe LV. 
FIGS. PAGE 
1, 2. Gervillia dentata Krauss . : 4 : : E ; : =< 


Sundays River Beds, Uitenhage Series, Cape Province. 


1. Right valve from the Sundays River, collected by Dr. W. G. 
Atherstone, Albany Museum 827, reduced x 0:7 
approximately, seen in lateral view. 

2. The same, interior view. 


Ann. S. Afr. Mus., Vol. XX XI. 


Photo: J. V. L. R. 


LAMELLIBRANCHIA FROM THE UITENHAGE 


Gervillia dentata. 


SERIES. 


Plate LY. 


Neill & Co., Ltd. 


ee ee eee 


ERN ZULULAND. 


owenya and Msinyene Pan. 


7. TABLE sHowING DistTRIBUTION oF Species 1N Lower CreTAckous Deposits, Poncora River AreA, NortHERN ZULULAND. 


Mfongosi. Ponpola, Myean Lombagwenya and Mainyene Pan, mae 
nd. 
7u.)7Z1.\22.|23,)24.)20.|26.\27.\28,)29.)210,)211.|z12 \z13.|214,)z16.)z17.\z19.)220,| 221, M1. | M3.) U1. | 12.) 03.|L3t.| 14. |05. | u6.|o7. | os. | 9. joan.|c12,.n141z04.|290, 
LAMELLIDRANCHIA. 
(Gucullaen|(Cyphoxis) ocala) 2 . callie z ; 
4 (Mlegnevculiaea) spp. | : : 
Arigonarea ot. ligeriensis me 2 
GimmeiOeytien o 3 2 6 A F 
Gervillia dentata . : és 3 : Dab escn | eoee | een een | ese x x x x x x x x 
Jeognomonsp. . = - = - x| x x x 
Inoceramus concentricus . o 5 6 ? i) x 
Wl \concenricuavar-eubeulcatua) x : : 
Peeudaviculat africana... «|x F s 
Arcelie priitevites 7) ee 1 
Erogyraconica =. wee t x x | x 1 t z 
Lophadiluviang 2 9. 0. ee 1 x 
inion » = Nell |pcleslells: x : x 
» _(lotrigonia) haughloni. . - || Sail lds 
(Rutitrigonia) pongolensis . x 
WN (eicurolrsgonsa) BUasceken Korn tae 3 
= \(Steinmanella)kennigh 9s Nx * < x ralles 
»  (Sphenotrigonia) frommurzei Set | sed | set | Se sc | ese x 
»  (Scabrotrigonia)spp.. . . x| x S| sel | sel | sed | eset || sect} seal x |x |} sll Riel x Ba hee 
sen 33 Verickis x || = 
ear ( 3 Yoh brava. 
Neitheaquadricosfala =. . . . ! x || x ? 
y quinquecostata 
Pholatomyatiuyness : 
aape viens ls «Nt ae : alalls 1 
Goniomya ap... we x x 
Ventdllatheridge) = 9. : mlles ? z 
Protocardia cf. sphaeroidea . . al| cele c 
Cardiumrogersi =. 2. 0 x |l x 
Rint so 2 o 6 all [eae Palle “ 2] x 2 : : x alts 
“WRG oo a 8 « x : 5 
GASTROPODA. 
Turritelamanvanensia » - : “alls 
Avelanawiineneata «9 6 : : ’ 
Dicroloma sp. tes «on gee x| x a 


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DESCRIPTIONS or toe PALAKONTOLOGICAL MATERIAL 
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: eel IV, containing : a 
10) A Critical Revision of the (pieces Perissodactyla of 
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( 393 ) 


10. A Critical Rersion of the Quaternary Perissodactyla of Southern 
Africa *.—By H. B. 8. Cooxz, M.A., D.Sc., F.G.S., F.R.Met.S., 
F.R.S.S. Afr. 


(With 30 Text-figures.) 


INTRODUCTION, 


THE study of fossil mammals in Southern Africa has been far from 
systematic and, with a few rare exceptions, writers have been more 
concerned with placing new species on record than with studying the 
fauna and revising our knowledge in the light of later discoveries. 
The first fossil mammal known to have been found in this region is 
the giant “Bubalus” bainw, whose horns and damaged skull were 
recovered in 1839 by the remarkable civil engineer and naturalist 
Andrew Geddes Bain from alluvial deposits of the Modder River, 
Orange Free State. The material was described only in 1891 by the 
British palaeontologist Seeley, and the next record of a fossil mammal 
appeared in 1906 when Dr. R. Beck described a mastodon tooth from 
the gravels of the Vaal River. In the following year another German 
scientist, Professor EH. Fraas, gave a further account of this tooth and 
commented on other remains from the gravels. Also in 1907, the 
celebrated American palaeontologist Professor W. B. Scott described 
a collection of fossil mammals from the coast of Zululand. Two 
years later Dr. Robert Broom made the first of his long series of 
contributions to mammalian palaeontology in South Africa with his 
descriptions of a new antelope from alluvial deposits at Caledon 
and of a new giant horse from a limestone fragment washed up 
on the beach near Maitland, in the south-western Cape. In 1913 
Broom described an assemblage of mammalian fossils from the 
thermal springs at Floris Bad, and in later years he described 
several new mammals from the Vaal River gravels and various 
open sites. 

In the past twenty-five years the initiative in the description of 
South African material has passed from the hands of outside experts 
like Seeley, Scott and Fraas to those of local workers such as Broom, 


* Manuscript submitted in January 1946. Appendix added in October 1950. 
VOL. XXXI, PART 4. 34 


FEB 7 195% 


394 Annals of the South African Museum. 


Haughton, Dart, van Hoepen, Middleton Shaw and others. During 
this period a considerable body of material has been collected, but 
much of it has remained largely unstudied and undescribed unless 
something obviously new was noticed by the individuals through 
whose hands it passed. Even then it has been for the most part only 
the new genera and species which were described, and the fauna as 
a whole has received little attention. Van Hoepen has large collections 
from his site at Cornelia which still await description, the South 
African Museum at Cape Town and the McGregor Museum at Kimber- 
ley have hundreds of specimens collected over a long period, and other 
museums have smaller quantities of undescribed material. Since its 
inception in 1934 the Archaeological Survey of the Union has acquired 
notable collections as a result of the activities of several collectors, 
and little of this material has been described or considered as a whole. 

During the years 1935-36 a joint survey of the Vaal River basin was 
carried out by Messrs. P. G. Séhnge and D. J. L. Visser of the Union 
Geological Survey, and Professor C. van Riet Lowe, Director of the 
Archaeological Survey, and during the survey much fossil material 
was recovered from various horizons in the deposits. The results of 
the geological and archaeological investigations were published in 
1937, under the title “The Geology and Archaeology of the Vaal River 
Basin”’, as Memoir No. 35 of the Union Geological Survey, and it was 
intimated in the letter of transmittal of this Memoir that the fossil 
material collected would be described at a later date. Through the 
courtesy of the Director of the Geological Survey all this material 
was placed in the hands of the present writer for examination and 
report.* 

It soon became apparent that a description merely of the material 
comprising this collection would be of little value, since it would not 
include all the species recorded from the deposits, and also because 
there occur in Pleistocene deposits other than those of the Vaal River 
basin many species which are likely at any time to be found within 
this area. For example, a tooth found at Christiana by a student and 
brought to the writer while this account was in course of preparation 
has been identified as belonging to a species hitherto recorded only 
at Cornelia in the Orange Free State. It was also only too obvious 
that the study of our fossil mammals had been far from systematic 
and that, with the notable exceptions of Haughton and Shaw, writers 
had been concerned more with placing new fossils on record than with 
comparative studies and revision of our knowledge in the light of 

* See Appendix. 


A Critical Revision of the Quaternary Perissodactyla. 395 


further discoveries. There exists a very considerable confusion of 
nomenclature and a multiplicity of specific names which renders the 
task of description of additional material virtually impossible unless it 
is accompanied by an amount of concurrent specific revision which 
would obscure the value of any account of the faunal assemblages. 
Indeed, it appears that until the material already described has been 
reviewed and new assessments made of the described species, it is 
of little value to proceed with the many other problems which our 
fauna presents. 

In the course of his investigations on the cave deposits, Broom has 
to a certain extent reviewed and revised the Primates, Insectivora, 
Rodentia and Carnivora. Furthermore, these orders furnish the bulk 
of the cave fauna, and are virtually unrepresented in the material 
from open sites. Most of the fossils recovered from the Vaal River 
basin, surface deposits and other open sites belong to the Perisso- 
dactyla, Proboscidea and Artiodactyla. It has accordingly been 
decided that before the undescribed material can profitably be dealt 
with, the described species of each of these three important orders 
of mammals in Southern Africa must be critically reviewed. The 
present paper considers the first, and perhaps the most important, of 
these orders.* 

As much as possible of the material from the Vaal River basin and 
elsewhere in the possession of the various museums in Southern Africa 
has been obtained on loan and examined in addition to the large 
collection in the Archaeological Survey. Dr. Broom has also been 
kind enough to make his material available to the writer. With the 
exception of those specimens which are in other countries, the type 
specimens of every species have been studied and are figured in the 
present account. Many of these figures have been drawn by the 
writer from the original specimens where the published figures are 
considered inadequate or unsuitable; others are reproductions of the 
original figures. As far as is possible new fossil material has deliber- 
ately not been introduced in this paper, as the purpose is to revise the 
specific descriptions from type material or from such other specimens 
as can with reasonable certainty be identified with the types. These 
descriptions are for the most part new, and are based on a fresh 
assessment of the original specimens interpreted in relation to the 
wider assortment of material now available and considered against 
a background of comparative researches upon the characteristics and 
variability of related living forms. It is hoped that this revision and 

* See Appendix. 


3 96 Annals of the South African Museum. 


correlation of scattered data may help to place our knowledge of these 
fossil mammals in Southern Africa on a firmer basis than has hitherto 
been the case, and serve to evaluate the characters of the revised 
species in a form which may facilitate future identification and 
comparison. 


MATERIAL. 


As a result of the conditions which prevailed in Southern Africa in 
the Quaternary, the fossil remains which have survived are mainly 
teeth, though skull fragments, loose bones and horn cores are some- 
times found under suitable circumstances. Almost all the described 
fossil mammalian species from this region have been named on teeth, 
and in the present account the descriptions of species generally give 
only their dental characters. Complete skulls are so rare that they 
need not be considered, and the identification of isolated bones is not 
yet possible, largely owing to the complete absence of whole or even 
partial fossil skeletons. 

The majority of the published descriptions have been rather un- 
systematic and often scanty. It has therefore been considered 
generally advisable not to quote the original account but to use it and 
the specimens in the preparation of a new description. Where 
quotations are given, however, the quoted material is indicated by 
the usual signs. The type specimens of each extinct species are 
figured and, in the descriptions given in the text, it has accordingly 
been possible to give an account of the features of the species which 
may be used for identification rather than a mere list of the character- 
istics of the particular specimen or specimens. An attempt has also 
been made here to give a definition of the generic characters of any 
extinct genus which has not been defined by its founder separately 
from the description of the genotype species. Where the genus is 
represented only by a single species, or perhaps by two species, this 
generic definition obviously may require considerable future revision, 
but some care has been taken to select as generic characters only the 
most outstanding features which differentiate the material from 
related types. 

The synonymy given for each species is as full as possible and, it is 
hoped, includes reference to all the specific designations given to fossil 
representatives of each described species. In the case of living 
species, however, the synonymy gives the reference to the type 
description only, and the further synonymy of the living forms can 


- A Critical Revision of the Quaternary Perissodactyla 397 


be found in the ordinary zoological sources: * the names applied to 
petrified specimens now assigned to these living forms are, however, 
given as fully as possible. The accepted specific names of all species 
living within historic times are followed by an asterisk, to avoid their 
confusion with species extinct prior to historic times and known only 
as fossils. 

In the descriptions of dental characters the nomenclature of the 
Cope-Osborn scheme has been followed as far as possible, in accordance 
with the generally accepted information available. Since there 
appears, however, to be some lack of uniformity in the conclusions 
regarding the homologies of the cusps, and there is occasionally some 
doubt regarding the application of certain terms, the terms applied 
in the present account are shown in a diagram of a typical member of 
each family described, or are clearly defined in the text. 

For convenience and brevity the customary abbreviations are used 
to denote molar, premolar, canine and incisor teeth. Milk teeth are, 
on the whole, of little value for specific identification owing to their 
considerable variability and the uncertainty of the relationship 
between their characters and those of the permanent dentition. 
They are accordingly considered only when absolutely necessary. 

In some cases a species is regarded as unrecognisable owing to the 
inadequacy of the material upon which it is founded or defined, and 
thus becomes a species insuff. descr. aut inqguirenda. It may neverthe- 
less be the case that a specimen which has been regarded as in itself 
inadequate for the creation of a species may be capable of reference 
to more adequate material. 

The numbers assigned by the various museums to the type and other 
specimens mentioned in this text are given wherever possible, together 
with an abbreviation indicating the museum concerned, viz.: 


Arch. Sur. Archaeological Survey, University of the Witwaters- 
rand, Johannesburg. 

Dept. Anat. Department of Anatomy, Medical School, University 
of the Witwatersrand, Johannesburg. 


M.M.K. | McGregor Memorial Museum, Kimberley. 
Nas. Mus. Nasionale Museum, Bloemfontein. 

S.A. Mus. South African Museum, Cape Town. 

Tvl. Mus. Transvaal Museum, Pretoria. 


* The most up-to-date synonymy is that given in “A Checklist of African 
Mammals’’, by G. M. Allen, Bull. Mus. Comp. Zool. Harvard, vol. Ixxxii, 1939. 


398 Annals of the South African Museum. 


ORDER PERISSODACTYLA. 
THE RHINOCEROSES. 


Amongst the fossil mammalia, the rhinoceroses are only very 
scantily represented in Southern Africa. Two supposedly extinct 
forms have been mentioned, each only from a single site, but petrified 
specimens indistinguishable from the two living species have been 
recovered from various superficial deposits. These latter specimens 
are probably not of any very great antiquity, but rhinoceros species are 
in any case not subject to rapid changes. 

The two living forms belong to different genera, and both their 
skulls and their teeth are quite distinct. The square-lipped or white 
rhinoceros is quite considerably larger than the hook-lipped or black 
rhinoceros, as can be seen from the drawings of their respective skulls 
(fig. 1). The lower jaws are sharply distinguished, that of the black 
rhinoceros having a deep compressed symphysis as compared with the 
depressed and rather spatulate symphysial region of the mandible in 
the white rhinoceros. The horns have been found isolated and again 
differ widely in form. Incisor and canine teeth are rudimentary or 
absent in both species. 

The cheek teeth in the rhinoceroses comprise four premolars and 
three molars arranged in a continuous series and having essentially 
the same structure, though the first premolar is considerably more 
simplified and is shed early. The lower third molar is also simple, 
and does not possess the third lobe so characteristic of the horses and 
most artiodactyls. The premolars are somewhat smaller than the 
true molars, the second premolar and first premolar particularly being 
smaller than the more uniform succeeding teeth. Structurally the 
teeth differ from those of the horse in being rather low crowned and 
in possessing strong, distinct roots, but their essential composition is 
similar to that of the equine cheek teeth. The normal order of 
eruption of the permanent dentition appears to be M!, Pm1, Pm?, M2, 
Pm’, Pm, and lastly M3, and is thus somewhat different from that 
of the horses. 

The cheek teeth of the rhinoceroses are lophodont in form, 7.e. the 
rows of cusps tend to become fused into ridges. In the upper teeth 
the two main outer cusps form a ridge known as the ectoloph, two 
anterior cusps form the protoloph and two posterior ones the metaloph. 
In the lower teeth three triangularly arranged cusps unite to form a 
crescentic metalophid, and posterior to this two cusps form an arcuate 


A Critical Revision of the Quaternary Perissodactyla. 399 


oO (0 20 30 40 50 690 


Centimet re Seale 


Fig. 1. 
Above: Lateral view of skull of Ceratotheriwm simum* (Burchell) and plan view 
of spatulate symphysial region of the lower jaw. (After Sclater.) 
Below: Lateral view of skull of Diceros bicornis* (Linnaeus) and plan view of the 
compressed symphysial region of the lower jaw. (After Owen.) 


400 Annals of the South African Museum. 


hypolophid ridge. With wear the enamel is rapidly removed from the 
top of these ridges and an area of dentine surrounded by enamel is 
exposed. This can be clearly seen in fig. 2, in which typical upper and 


prefossette 


Metafiexid Entoflexid 


Fie. 2.—Molar elements (following Osborn) of the upper and lower cheek - 
teeth of the Rhinoceros group. 
Abbreviations. 
Upper teeth: pas, parastyle; pa, paracone; me, metacone; hy, hypocone; 
pr, protocone; pel, protoconule; mcl, metaconule. 
Lower teeth: prd, protoconid; hyd, hypoconid; pad, parastylid; med, meta- 
conid; end, entaconid. (Original.) 


lower first molars are shown indicating the nomenclature used for the 
cusps, folds and ridges (following Osborn). The valley between the 
protoloph and metaloph appears to have received no name, and is 
here termed the medivallum by analogy with the corresponding 
valley in horse teeth. For the two inlets in the lower teeth the terms 
““metaflexid”’ and “‘entoflexid” are here suggested for convenience 


A Critical Revision of the Quaternary Perissodactyla. 401 


in description, as the same terms have been proposed by Stirton (1941) 
in the lower teeth of the horses. 

Owing largely to a lack of sufficient material, it has unfortunately 
not been possible to gain any reliable idea as to the constancy or 
variability of the tooth characters within the series in the rhinoceroses. 
From the limited material examined, however, it does appear that 
while the essential structures are reasonably constant, the effect of 
attrition alters the pattern of the grinding surface to such a degree 
that identification may be made most difficult. As wear proceeds, the 
ridges widen and obliterate the intervening valleys, at first fairly 
slowly, but afterwards very rapidly, until ultimately a uniform tract 
of dentine may be produced. The crochet, antecrochet and crista 
which project into the medivallum are generally more prominent in 
early wear, and are reduced in size as this valley is narrowed. In some 
species the crochet and crista may unite and isolate the medifossette 
as an accessory valley, leaving the prefossette as the terminal portion 
of the medivallum. The postfossette may also become isolated by 
closure of the posterior enamel border. In the lower teeth the chief 
effect of attrition is to reduce the size of the two flexids, the metaflexid 
in particular tending to disappear with wear. Fusion of the meta- 
conid and entaconid may also lead to the complete isolation of the 
entoflexid as an accessory valley. 


Famity RHINOCEROTIDAE. 
Genus Dicrros Gray 1821. 
Genotype: Rhinoceros bicornis* Linnaeus. 
Diceros bicornis® (Linnaeus). 


Rhinoceros bicornis®* Linnaeus 1758. Syst. Nat. Ed. (10), i, p. 56. 
Opsiceros simplicidens (pars) Scott 1907. 3rd Rep. Geol. Surv. Natal 
and Zululand, pp. 258-259, pl. xvu, figs. 4, 5. 
Diceros whitei (pars) Chubb 1907. Geol. Mag., V, vol. iv, pp. 447-448. 
The horns of the black rhinoceros are almost invariably two in 
number, but exceptionally as many as five have been recorded. The 
anterior horn has a height of about forty-five to sixty centimetres on 
the average and has a basal diameter fifteen to twenty-five centi- 
metres. The rear horn is about one-third to one-half the length of 
the anterior one and has a diameter only a little less than its height. 
The record horn lengths are about double the average figures. Both 
horns are rather blunt and curve very slightly posteriorly. 
In the upper jaw the first premolar is very small and exhibits no 


402 Annals of the South African Museum. 


structures which can be said to be recognisable as persistent. The 
third molar is triangular in form, the metaloph, being reduced to a 
small posterior prominence only, and this tooth is very variable in 
pattern in wear and is of little value for specific identification. The 
remaining three premolar and two molar teeth are generally more 
consistent, and are essentially similar in structure, though the pre- 
molars differ slightly from the molars. In the premolars the anterior 
wall of the protoloph is fairly straight and makes an angle of about 
75° with the ectoloph, whereas in the molars the protoloph initially 
makes almost a right angle with the ectoloph, and then curves some- 
what posteriorly. The protoloph and metaloph are roughly parallel or 
slightly divergent and with the ectoloph give the appearance of the 
Greek letter 7. The ectoloph itself is not straight, but has an outer 
wall incurved or grooved between the paracone and metacone and also 
has a shallow groove behind the parastyle. The parastyle itself is 
commonly anteriorly grooved and projects very little in front of the 
protoloph. The antecrochet is apparently absent, and the crista is 
very small and disappears rapidly with attrition. A crochet is always 
present in the earlier stages of wear, and in the normal dentition 
increases progressively in size from the second premolar to the second 
molar. It tends to become rounded with increased wear, and may 
disappear completely before the medivallum is obscured. In no case 
has isolation of the medifossette been observed in this species except in 
the third molar. The postfossette is somewhat obliquely V-shaped, 
tending to be U-shaped with wear as a result of the expansion of the 
hypocone lobe of the metaloph, and then becomes isolated as an oval 
valley. The dimensions vary considerably with wear, the breadth 
across the grinding surface increasing as attrition proceeds. The 
height above the basal cingulum increases progressively with the 
successive teeth, and a typical second molar in early wear has a height 
of about 50-55 mm. The breadth at the base of the second molar is 
about 60 mm., but in normal wear the grinding surface measures only 
some 45 mm. transversely. The size and characters can be seen from 
the scale drawings in fig. 3. Two typical upper dentitions are shown, 
one in fairly early wear, the other well worn and lacking the first 
premolar. 

The lower teeth have little to distinguish them from the very 
generalised form of most rhinoceros teeth. The first premolar is 
greatly simplified in form, but the remaining teeth, including the 
third molar, are similar in structure. The anterior and antero- 
external walls of the metalophid are markedly flattened, and make an 


A Critical Revision of the Quaternary Perissodactyla. 403 


< 


Fic. 3.—Two left upper dentitions (A and B) of Diceros bicornis® (Lin- 
naeus) and a right lower dentition (C) of the individual B. One-half 
natural size. (Original.) 


404 Annals of the South African Museum. 


angle with each other of about 100° ora little more. A fairly marked 
groove separates the outer wall of the metalophid from the curved 
hypolophid walJl. The inner walls of the metaconid and entaconid are 
somewhat flattened. The metaflexid is a good deal smaller than the 
entoflexid and is rapidly reduced to a shallow V-shaped notch. The 
height of a normal second molar is about 50mm. The lower teeth of 
a typical specimen are shown in fig. 3, and belong to the same indi- 
vidual as the upper dentition figured immediately above it. 


Referred Material. 


Apart from the petrified specimens from superficial deposits in 
various parts of Southern Africa which obviously belong to this species, 
two of the teeth from Zululand which Scott (1907) very tentatively 
referred to his species Opsiceros simplicidens do not appear to warrant 
distinction from the living Diceros bicornis*. The type LM? of Scott’s 
species manifestly is not that of D. bicornis*, but the two heavily worn 
teeth (M! and Pm+*) do not differ appreciably from correspondingly 
worn teeth in old individuals of the living black rhinoceros. Scott 
himself realised the close similarity, and suggested that these two 
teeth did not actually belong to his new species. The specimens 
themselves have not been seen by the present writer, but natural size 
photographs kindly supplied to the writer by Professor Scott, together 
with the admirable description, form an adequate basis for the 
conclusion reached above. 


Genus CERATOTHERIUM Gray 1868. 
Genotype: Rhinoceros simus* Burchell. 


Ceratothervum simum* (Burchell). 


Rhinoceros stmus* Burchell 1817. Bull. Sci. Soc. Phil. Paris, F. 1, 2, 
105 Sil 

Opsiceros simplicidens Scott 1907. 3rd Rep. Geol. Surv. Natal and 
Zululand, pp. 257-258, pl. xvii, fig. 3. 

Rhinoceros scotti Hopwood 1926. Occ. Papers No. 2, Geol. Survey, 
Uganda, pp. 16-17, fig. 3. 


The white rhinoceros possesses a long and rather slender anterior 
horn which attains a height of about ninety centimetres and, exception- 
ally,as much as a hundred and fifty centimetres.* The second or rear 


* Southern race: 624 inches. Rowland Ward’s Records of Big Game, 9th ed., 
1928, p. 446. 


A Critical Revision of the Quaternary Perissodactyla. 405 


horn 1s small and does not usually attain a height exceeding twenty-five 
centimetres. The anterior horn is normally more slender than that of 
the black rhinoceros. 

In the upper jaw the first premolar is small and is shed early, but 
the other premolars differ notably from the molars, for in the former 
the medivallum becomes rapidly enclosed by fusion of the proto- 
cone and hypocone, the form of the metaloph being consequently 
also affected. From Diceros bicornis* they differ most markedly in the 
arcuate, posteriorly curving protoloph, and in the early fusion of the 
well-developed crista with the crochet to form an isolated medi- 
fossette. Due to the posterior curving of both protoloph and metaloph 
the m-like shape of the teeth is very distorted and not nearly as 
noticeable as in D. bicornis*. The form of the ectoloph is also 
different, being rather more undulate in Ceratotherium simum* with 
a distinct outward bulge at the paracone. The parastyle is more 
pointed and is not apparently grooved, but there is a groove immedi- 
ately behind it. The postfossette becomes isolated with wear as a 
result of closure of the posterior enamel. As in D. bicornis* this 
isolation of the postfossette is not a constant feature and is generally 
less marked in the molars than in the premolars. The teeth are 
higher crowned than those of the black rhinoceros, a typical second 
molar in normal wear measuring about 75 mm. above the indistinct 
cingulum. Fig. 4 shows (half natural size) two upper dentitions, one 
in very early wear with the third molar only just erupting and the 
fourth premolar coming into use, the second dentition being in a more 
advanced state of attrition. 

The lower teeth are somewhat difficult to distinguish from those of 
the black rhinoceros. The anterior and antero-external walls of the 
metalophid make an angle with each other close to 90° instead of the 
obtuse angle found in D. bicornis*. The metaflexid appears to be 
more persistent in the white rhinoceros than in the black, and the 
enclosure of both metaflexid and entoflexid with advanced wear is a 
common feature. The height of a typical second molar above the 
cingulum is about 60mm. The lower dentition of the same individual 
as the more worn upper dentition figured is shown in fig. 4. 


Referred Material. 


Petrified specimens of this species have been found in various 
superficial deposits in the coastal region and in the interior. A portion 
of an anterior horn is also recorded from a cave deposit near Kuruman, 
Cape Province (Malan and Cooke, 1941). 


406 Annals of the South African Museum. 


Fie. 4.—Two left upper dentitions (A and B) of Ceratotherium simum* 
(Burchell) and a lower right dentition (C) of the individual B. One-half 
natural size. (Original.) 


A Critical Remsion of the Quaternary Perissodactyla. 407 


W. B. Scott reported in 1907 the discovery of a supposedly extinct 
species in fossiliferous marine clays from the Zululand coast, and he 
named this species Opsiceros simplicidens, with an unworn upper left 
second molar as the type. Scott compares this tooth with the 
corresponding one of Diceros bicornis*, and states that the differences 
are “clearly of specific value”. The distinctive characters are stated 
‘to lie, enter alia, in the much stronger recurving of the protoloph and 
in the much better development of the crista, which “fuses with the 
anticrochet so as to enclose a small and apparently shallow fossette’’. 
(Scott here used the term ‘anticrochet’ in error for crochet, as his 
figure shows, and in this follows an error previously made by Osborn 
in describing the Perissodactyla of White River (Scott and Osborn, 
1890).) The characters which Scott used to differentiate the species 
from D. bicornis* are exactly those which distinguish the white 
rhinoceros. A skull of the latter species in the South African Museum 
possesses a second molar in much the same state of development, 
and the dimensions and appearance of this tooth correspond very 
closely to the data and figure furnished by Scott. There can thus be 
little doubt that Opsiceros simplicidens is a synonym of Ceratothervum 
simum*, a fact which Scott would undoubtedly have realised had 
comparative material of this rather rare species been available to him. 

In 1926 Hopwood recorded an upper left second molar from the 
Kaiso beds of Uganda and assigned this specimen to Scott’s species. 
He also pointed out that the name R. simplicidens was preoccupied 
and proposed Rhinoceros scotti as a substitute. There seems no doubt 
of the correctness of the reference of this specimen, and equally there 
is little doubt of its similarity to teeth of the living white rhinoceros. 
Rhinoceros scotti is thus also apparently a synonym of Ceratotherium 
sumum*. 


DISCARDED SPECIES. 
Diceros white Chubb. 


Diceros whitei Chubb 1907. Geol. Mag., V, vol. iv, pp. 447-448. 
Diceros whitei Hopwood 1928. Rhodesian Man and Associated 
Remains. 


A supposedly new species of rhinoceros was described very briefly 
by E. C. Chubb in 1907 in a “List of Vertebrate Remains” from the 
Broken Hill Cave. It was founded on two limb bones (a right tibia 
and a right humerus) which had been excavated by Mr. Franklin 


408 Annals of the South African Museum. 


White and presented by him to the Rhodesian Museum. This species 
was also mentioned by A. T. Hopwood in the British Museum memoir 
on Rhodesian Man, and is there said to be “closely allied to D. stmus”’. 
In view of the uncertainty of the generic position of the species whitet, 
the material was obtained on loan from the Rhodesian Museum, 
Bulawayo, and permission was obtained from Mr. Chubb to amplify 
his preliminary description and to figure the specimens. In his brief 
account Chubb remarks on the scantiness of the comparative material 
available to him, and this lack and the seeming association of the two 
bones appears to have resulted in an error in the distinction of the 
material. The tibia is certainly that of a rhinoceros, though com- 
parison with recent skeletons shows no notable differences in size or 
in other characters from the corresponding bone in the living Diceros 
bicornis*. The humerus, however, differs very considerably from 
both the living rhinoceroses, and it would appear that it is an artio- 
dactyl and not a perissodactyl humerus, the differences formerly 
regarded by Chubb as of specific distinctness being actually too great 
for that possibility to be upheld. The compressed narrow olecranon 
fossa is a normal artiodactyl feature unlikely to occur in a rhinoceros, 
and the deltoid ridge and deltoid tuberosity are also much more 
artiodactyl than perissodactyl. With these views Mr. Chubb now 
expresses his agreement. 

On comparison with various living artiodactyls, the closest resem- 
blance is found between the fossil humerus and that of the living Cape 
Buffalo. There is no great difference in length, but the fossil bone is 
somewhat more massive, with the attendant minor modifications 
consequent upon its greater weight-supporting requirements. Other- 
wise, however, there is a very close agreement in every character, 
and it seems highly probable that the fossil humerus belongs to a 
member of the Buffalo group. It may possibly belong to the extinct 
“ Bubalus”’ baini Seeley, or to “‘Bubalus” andersoni Scott. 

The species Diceros whiter appears, therefore, to have been founded 
on a humerus which is not that of a rhinoceros and on a tibia which 
does not warrant distinction from the living D. bicornis*, so that 
D. whitei must be regarded as incorrectly founded. 


THE Horsks. 


There have been described at various times from Southern Africa 
more than twenty-five species belonging to this family, some based on 
upper and some on lower teeth, but of these not more than half can be 


A Critical Revision of the Quaternary Perissodactyla. 409 


regarded as valid. The position was first reviewed by Haughton in 
1931 when the twenty then existing species were reduced to eleven, 
and in general the present writer is in agreement with these con- 
clusions. Haughton divided the members ascribed to the genus 
Equus into two groups, which he terms the “quagga”’ group and the 
“zebra” group on the parallelism of certain characters with those in 
the teeth of the two living species, the bontequagga and the mountain 
zebra. Unfortunately these characters in the recently extinct true 
quagga are very different from those in the living bontequagga and 
the two group terms must therefore be abandoned. 

The relationships indicated by Haughton’s work are of great 
interest and, in order the better to appreciate the definitive characters 
of the teeth in the extinct forms, the writer has carried out an extensive 
examination of skulls and teeth of the living forms and of the recently 
extinct true quagga. As a result of this work it is possible to dis- 
tinguish on dental characters from this material three undoubted 
species: Hquus zebra*, the living mountain zebra, Equus quagga*, 
the recently extinct true quagga, and Equus burchellii*, the living 
bontequagga or Burchell’s zebra. Since zoologists have been greatly 
at variance on the status of these forms, and since all three species 
occur in the fossil state, the results of the investigation have already 
been considered fairly fully (Cooke, 1943). These observations also 
throw some light on the morphological characters and variations 
encountered in equine species and are of great value in considering 
the fossil finds. To some extent they repeat and amplify the work 
of Gidley (1901), and in the present examination a general agreement 
was found with the conclusions outlined by him. 

For convenience of reference the nomenclature of the important 
elements of the molar teeth of the Equidae (following Osborn) is given 
here in diagrammatic form (fig. 5). The specimens figured are upper 
and lower fourth premolars, and show the appearance of the cusps 
on the unworn crowns and the enamel patterns of the teeth in normal 
wear. The two enamel islands in the upper cheek teeth have long 
been known as the pre- and postfossettes, but the partial islands, or 
inlets, in the lower teeth have until recently received no name. The 
terms advocated by Stirton (1941) are used here. They are re- 
spectively “‘metaflexid’”’ for the anterior and “entoflexid” for the 
posterior partial islands of the lower cheek teeth. (These terms have 
already been suggested for the analogous parts of the rhinoceros teeth.) 
It is also proposed here to call the posterior groove which lies between 
the hypocone and the hypostyle in the upper teeth the “hypoglyph”, 

VOL, XxXxt, PART 4, 3D 


410 Annals of the South African Museum. 


and the groove anterior to the protocone the “protoglyph”. These 
terms are preferred to ‘“‘hypoconal groove” and “preprotoconal 
groove’ used by Stirton. 


Ectoloph 


a 
%, 
Medivallum 
pli pretossette 
pli postfossette 
protoloph 


pli hypostyle 
pli protoconule Hypoglyph 


pli caballin 


Metaflexid Entorlent. 


Fig. 5.—Molar elements (following Osborn) of the upper and lower cheek 


ARG RCDATIONS teeth of the Horse group. 


Upper teeth: pas, parastyle; mss, mesostyle; mts, metastyle; pa, paracone; 
me, metacone; hy, hypocone; pr, protocone; pel, protoconule; mcl, metaconule; 
hys, hypostyle. 

Lower teeth: prd, protoconid; hyd, hypoconid; med, metaconid; end, enta- 
conid; pad, parastylid; msd, metastylid; hld, hypoconnlid; ec.sd., ectostylid 
(fold or ridge); esd, entostylid. (Original.) 


Distinction between Zebra, Quagga and Bontequagga. 


The writer has examined a large number of skulls of these three 
species in the past few years, though skulls of the recently extinct 


~ Sinai anal 


A Critical Revision of the Quaternary Perissodactyla. 411 


quagga are rare and difficult to obtain. Owen (1869) figured one in 
a little known paper, there is a cast in the Transvaal Museum, one 
skull in the Kingwilliamstown Museum and several in the McGregor 
Museum, Kimberley. Based on an examination of this material the 
chief distinguishing features have been analysed (Cooke, 1943). 

From the point of view of classification it would appear that the 
three species are quite distinct. The bontequagga or Burchell’s zebra, 
while showing skull characters intermediate between the other two 
forms, is sharply distinguished in dental characters. There can thus 
be no doubt that the bontequagga is very distinct from the true zebra 
despite the many similarities of form and colouring which have caused 
much dispute amongst zoologists with respect to the relationship of 
the two forms. It differs to an equal degree from the true quagga, 
and can under no circumstances be regarded as a variety of this form. 
The specific designation guagga* must therefore be restricted to the 
historically extinct true quagga, and the bontequagga or Burchell’s 
zebra must receive the full specific name burchellii*, to which may be 
appended, if it is considered necessary, the varietal names wahlbergi*, 
transvaalensis*, etc., though these varieties cannot be distinguished 
on skeletal or dental grounds as far as the present writer is aware. 


The Degree of Constancy and Range of Variability of the 
Tooth Characters. 


Aside from the examination of a large number of skulls of the 
zebrine group of horses to determine their distinctive features, an 
attempt has been made to estimate the range of variation found 
within each species and hence to estimate the value of each possible 
factor in specific determination. A number of skulls of Equus 
caballus* and Equus asinus* have also been used for this purpose, and 
reference has been made in addition to many published figures of 
equine dentitions to ascertain how widely the generalised conclusions 
may be applied. The factors appear from this examination to follow 
certain definite trends, a knowledge of which greatly enhances the 
value of the specific identification of individual finds of isolated teeth. 
It must be stated however that, despite this knowledge, a fossil species 
which is named purely on dental characters may well be a “form” 
species only, as there is a certain amount of overlap in the extreme 
variations of certain species. The first upper true molar of Hquus 
quagga*, for instance, may be almost identical with that of Equus 
zebra* under certain conditions of variation. 


412 Annals of the South African Museum. 


One of the difficulties which besets the worker on the fossil Equidae 
is the fact that almost all the finds consist only of isolated teeth. 
Complete dentitions are very rare and are consequently of immense 
value. It is of the utmost importance therefore to ascertain correctly 
the position occupied by the isolated teeth in the former jaw in order 
correctly to evaluate the determinative characters. By far the most 
satisfactory method is that of direct comparison with a known complete 
dentition. The angle of wear and degree of antero-posterior curvature 
are the chief guides in the estimation as to whether a particular tooth 
is, for example, a fourth premolar or a first molar. Fig. 6 gives an 
indication of the general shape of the teeth in Equus burchellii*, and 
figs. 6 and 19 show the form in some large extinct equines. 

The second premolar is at once distinguishable by being rather 
pointed anteriorly, and the third molar shows a posterior tapering. 
The last erupted tooth of the series at any given age shows this posterior 
taper, however, so that in early wear such a tooth might possibly be a 
second or even a first molar. The rather sharp curvature of the true 
third molar should, however, facilitate the distinction of this tooth. 
In the lower teeth the premolars typically have a larger entoflexid 
than the molars, and the outer groove between hypoconid and proto- 
conid extends further towards the inner groove between metaconid 
and metastylid in the true molars than it does in the premolars. In 
the upper teeth the first true molar normally has a rather narrow 
mesotyle, but otherwise the degree of curvature and angle of wear 
form the only guide to the position of the tooth in the series. 

The factors which affect the characters of each cheek tooth in an 
individual of a species are: (A) The position of the tooth in the series, 
(B) the degree of attrition, and (C) variability within the species. The 
factor of sex does not appear to exert any appreciable influence on the 
cheek teeth, though the canines, which are usually prominent in the 
male, are vestigial or absent in the female. The generalised con- 
clusions regarding the three main factors given above may therefore 
be discussed in turn. 


A. THE PosIrion oF THE TOOTH IN THE SERIES. 


(a) In the Upper Teeth. 


1. The anterior and posterior teeth (second premolar and third 
molar) show decidedly different forms of enamel pattern due to 
their tapering nature and are very variable in character. They are 
of the least possible value in specific determination, as the prefossette 


Above: 


Below: 


Fic. 6. 
Lateral view of skull of Equus burchellii* (Gray) showing the shapes of 
the cheek teeth. (Original.) 
Lateral view of fragmentary remains of skulls of large extinct equines; 
Maxilla of Hquus fowleri Wells, and mandible of Equus plicatus (van 
Hoepen). (Original.) 


~—eow @ 2 2 @ =a 
~~. 


414 Annals of the South African Museum. 


in the second premolar and the postfossette in the third molar are 
subject to considerable distortion. 

2. In transverse breadth, measured from mesostyle to protocone, 
the third and fourth premolars are approximately equal, and the first 
and second molars usually slightly narrower (by perhaps 5-10 per cent.) 
than the preceding teeth. The second premolar and third molar are 
always narrower than the other premolar or molar teeth respectively, 
but bear no constant ratio of relative size. 

3. The fourth premolar presents the most complex and the first 
molar the simplest enamel folding.* 

4. The protocone is shortest (antero-posteriorly) in the second 
premolar and longest in the third molar. The length of the protocone 
either increases progressively in each succeeding tooth of the series 
or else the first molar has a smaller protocone than the fourth premolar, 
and the second molar has one about the same length as in the fourth 
premolar. 

5. The mesostyle is generally reduced in width in the molars as 
compared with the premolars. 

6. The pli-protoloph and pli-hypostyle may be lacking in the first 
molar though present in the remaining teeth. 

7. The pli-caballin may be present in the premolars yet absent in 
the molars, and may occur in the fourth premolar only. 

8. The hypoglyph is often absent in the third molar. 


(b) In the Lower Teeth. 


1. As in the upper teeth, the second premolar and third molar are 
of sub-triangular form and show departures from the normal enamel 
pattern. The changes are not always so great as to prevent specific 
identification, but they are the least useful teeth of the series for this 
purpose. 

2. In transverse breadth the premolars are wider than the molars 
by about 5-10 per cent. The second premolar may, however, be as 
narrow as the first molar. The third molar bears no reliable ratio of 
width to the other molars, and in the second molar the posterior 
breadth is less than the anterior due to the reduction in size of the 
hypoconid. 


* ‘The normal order of eruption of the cheek teeth is M2, M2, Pm?, Pm’, Pm4, and 
lastly M%, though the order of the last two may exceptionally be reversed. In a 
normal dentition, therefore, the first molar in addition to possessing an initially less 
complex character is also the most worn tooth. 


A Critical Revision of the Quaternary Perissodactyla. 415 


3. The entoflexid has a larger lobe in the premolars than in the 
molars, this being generally of greatest antero-posterior length in the 
second premolars. 

4. The outer groove between protoconid and hypoconid is deeper in 
the molars than in the premolars and sometimes meets the internal 
groove between metaconid and metastylid. 


(c) Relation between Upper and Lower Teeth. 


Of great interest and value is the hitherto apparently unrecognised 
fact that the breadth over the enamel in the upper teeth bears a fairly 
constant ratio to the breadth of the lower teeth over the enamel. This 
ratio is approximately 1: 0-6 for the third and fourth premolars to 
_1: 0-55 for the first and second molars, and is apparently subject to a 
variation of only about 5 per cent. 


B. Tut Decree or ATTRITION. 
I. Changes in Dimensions. 


The effects of the degree of wear (or age) on the cheek teeth have 
been well described by Gidley (1901), and the following selected 
quotations from his paper are illustrative of his findings in this 
regard. 


1. “When a molar or premolar tooth first comes into use, the face, 
as well as the sides of the crown, is completely covered with enamel 
which folds in and out, and (though somewhat hidden by cement) 
presents the same general appearance as that seen in the much more 
primitive forms Anchitherium and Mesohippus. Soon the enamel on 
the tips of the cones and along the ectoloph wears through, and small 
patches and ridges of dentine surrounded by a border of enamel are 
exposed. As the tooth is further worn away, these patches and 
irregular ridges broaden and rapidly lengthen until when about one- 
half to three-quarters of an inch of the crown has been worn away they 
have all become united by narrow isthmuses, and the fundamental 
tooth pattern of the horse is presented.” “From this point... 
the triturating surface presents a gradually less complex pattern of 
enamel folding as the tooth crown is worn away, until in the very 
much worn tooth the simplest pattern of enamel folding is presented” 
tee 0) , 

2. ! The antero-posterior diameter of the first premolar (p,)* 
remains about the same for the whole length of the crown, except 
that sometimes it narrows slightly near the roots.” 


* Actually the true second premolar according to present usage. 


416 Annals of the South African Museum. 


Fic. 7.—Sectioned right upper fourth premolar of adult male Equus 
caballus* Linnaeus, illustrating changes in enamel pattern which would 
result from various stages of attrition. Natural size. (Original.) 


3. ‘The antero-posterior diameter of the last molar (m3), however, 
is relatively small at first, and increases continually as the tooth is 
worn away.” 

4. “The antero-posterior diameter of the grinding surfaces of all 


A Critical Revision of the Quaternary Perissodactyla. 417 


the intermediate teeth are greater at the stage when the tooth has just 
fully come into use; ...from this point the antero-posterior 
diameter diminishes very rapidly for a short distance and then con- 
tinues to diminish more gradually to the roots of the tooth.” 

5. After the tooth has just come fully into use “the transverse 
diameters of p* to m? inclusive remain about the same, diminishing 
slightly near the roots; p* gradually diminishes while m? increases in 
transverse diameter as the crown wears away.” 

6. “The antero-posterior diameter of the protocone in all the teeth 
of the series remains the same for the whoie length of the crown.” 

7. “Owing to the very slight variation of the transverse diameters 
of the crowns of p* to m? inclusive . . . and to the great shortening of 
their antero-posterior diameters, the ratio of these diameters is very 
different in old and in young individuals of the same species. Thus 
in the little worn condition . . . the antero-posterior diameter is 
always greater than the transverse. As the crown wears away... 
a stage is reached where the two diameters are about equal, then, 
as the antero-posterior becomes still more shortened, the transverse 
exceeds it. In every series this variation in ratio seems always to be 
more advanced in m! and m?.” 


The present writer has not observed any conflict with these views. 
The transverse diameter (or “breadth’’) of the crown and the antero- 
posterior diameter (“‘length’’) of the protocone do, however, decrease 
to a very slight degree with wear, though their ratio remains constant. 
Gidley made no comments on the precise effects of wear on the com- 
plexity of the enamel folds, and the present writer has therefore 
attempted by means of serial sections of a number of teeth to ascertain 
which folds are most affected by wear and which least. As a result, 
the following generalisations, though based on comparatively little 
material, appear to be possible. 


il. Changes in Enamel Complexity. 
(a) In the Upper Teeth. 


1. The pli-protoloph may disappear completely when the tooth is as 
little as half worn, and is always the most affected of the fossette 
folds. 

2. The pli-hypostyle and pli-prefossette suffer rapid reduction in 
size but have not been observed actually to disappear. 

3. The secondary small plications in the regions of the pre- and 
postfossette folds proper disappear rapidly if they are of an angular 
or saw-tooth nature, but may survive a considerable degree of wear 
if they have the form of rounded loops. 


418 Annals of the South African Museum. 


4, The pli-protoconule and pli-postfossette are the least affected 
of the fossette folds. 

5. The hypoglyph becomes shallower with wear but does not 
normally disappear. 

6. The pli-caballin does not disappear with wear in the premolars 
unless initially very small, but may do so in the molars. 

7. The halves of the ectoloph become slightly flattened with 
advanced wear. 


(b) In the Lower Teeth. 


1. The folding of the entoflexid lobe may be very complex in the 
earliest stages of wear, but becomes rapidly simplified and does not 
normally survive the half-worn condition of the tooth. 


C. VARIABILITY WITHIN THE SPECIES. 
I. Transverse Diameter of the Crown. 


As Gidley has shown, the transverse diameters over the enamel of 
the teeth (excluding the second premolar and third molar) are subject 
only to relatively slight variation within a species. In a series of ten 
skulls of Equus caballus* ranging from a large draught horse to a small 
Texas pony, he notes a difference in the transverse diameter of the 
second molar of only 2-5 mm. or about 10 per cent. In the third 
premolar of the same individuals, however, his figures show a difference 
of 4-5 mm. or nearly 20 per cent., and this range of variation agrees 
with the present writer’s findings in the case of the zebrine group. 
In the many skulls of Equus burchellui*, Equus zebra* and Equus 
quagga* which have been examined, the extreme variations in trans- 
verse diameter amount to about 20 per cent. of the mean value, but 
at least 70 per cent. of the specimens in each species are very close 
indeed to this mean value. The transverse diameter of the cheek 
teeth is thus subject to a variation of about plus or minus 10 per cent. 
from a predominant mean value, and this value is therefore a useful 
characteristic of a species (except in the anterior and posterior teeth). 
The same relative figures apply to the lower teeth. 


II. Length and Form of the Protocone. 


(a) The form of the protocone, and particularly the relative anterior 
and posterior development or elongation, is a very constant character 
in a species. 


A Critical Revision of the Quaternary Perissodactyla. 419 


(0) The absolute length of the protocone is subject within a species 
to a variation of as much as 30 per cent. between corresponding teeth. 
In general, the ratio of the length of the protocone to the transverse 
diameter of the crown is not affected by wear, and this ratio (which we 
may term the “protocone ratio”) is not quite as variable as the 
absolute length of the protocone. In Equus caballus* the average 
value of the protocone ratio 


length of protocone 


transverse diameter of crown 


is 0-48 in the fourth premolar; in the corresponding tooth of Equus 
asinus* 0-41; Hquus burchellii* 0-41; Equus zebra* 0-40; Equus 
quagga* 0-44. Though subject to a variation of plus or minus 15 per 
cent., and thus having no precise significance since values would 
overlap considerably, the protocone index might be a useful indication 
of the relative development of the protocone in a species. It must be 
remembered, however, that in addition to variation between corre- 
sponding teeth, the value of the protocone ratio will change from the 
second premolar to the third molar. In Hquus caballus* average 
figures are p* 0:38, p? 0-45, p4 0-48, m! 0:47, m? 0-52, m® 0-62. 


Ill. The Enamel Folds. 


A wide variation is found in the degree of complexity of the enamel 
folding within a species, but the following generalisations appear from 
the writer’s investigations to be applicable to the upper teeth. 

1. The caballine fold is not infrequently absent in particular teeth 
of an individual of a species which normally shows a strong caballine 
fold. Even though in some species it is apparently never present, it 
has no precise value. 

2. The pli-protoloph is of very variable development and unless 
very strong is of no specific value. 

3. The degree of development of the pli-hypostyle and pli-prefossette, 
and the secondary plications of the latter, is variable and of slight 
value only. 

4, The pli-protoconule and pli-postfossette appear to be of very 
constant development and their presence or absence can be considered 
as of diagnostic importance. 

5. In some species the hypocone shows a consistent slight bulge into 
the medivallum, giving it the appearance of a partial isolation from 
the metaconule, and this character seems to be of some determinative 


value. 


420 Annals of the South African Museum. 


6. The flattening of the inner wall of the protoconule appears to be a 
character of minor diagnostic value in some species. 
7. The depth of the hypoglyph is a moderately constant feature. 


IV. The Ectoloph and Styles. 


1. Except in the second premolar and third molar, the nature of the 
halves of the ectoloph, more particularly the anterior one, is of fairly 
constant character, being either concave inwards as seen in Equus 
caballus* and Equus burchellii*, or else flattened or even convex as in 
Equus asinus*, Equus zebra* and Equus quagga*. 

2. The three styles, parastyle, mesostyle and metastyle, show quite 
a considerable range of variation in detail within a species in addition 
to the difference shown between premolars and molars. The characters 
of the metastyle are of no specific value, and neither the absolute size 
nor the grooving of the parastyle and mesostyle are entirely constant 
characters though they may be useful ones. The degree of isolation 
of the mesostyle and parastyle from the walls of the ectoloph or their 
easy confluence with them, however, appears to be a valuable specific 
character. Owing to the normal relative reduction in size of the styles 
in the true molars, their isolation is less apparent in these teeth than 
in the premolars. 


V. The Lower Teeth. 


To quote from Gidley, “the characters of the lower teeth are, in 
general, affected in the same way as the upper, and seem to be of even 
less value in determining the species”. With these conclusions the 
present writer is in agreement, in that the very simplicity of structure 
of the lower teeth deprived them of many of the characters which 
might be of value in making distinctions. The range of normal 
variation within a species thus permits of considerable overlap 
between what are actually different species (as is only too clearly 
seen in the zebrine group of horses), and thus absolute size must 
be the chief determinative factor. Other useful, though not entirely 
constant characters, are (a) the shape of the outer walls of proto- 
conid and hypoconid, (6) the shape and degree of separation of 
metastylid and metaconid, (c) the size and form of the metaflexid and 
entoflexid lobes, (d) the relative size of the stylids and conids, (e) the 
development of the ectostylid fold* in the hypoconid. 


* Called “pli-caballinid’’ by Stirton (1941). 


A Critical Revision of the Quaternary Perissodactyla. 421 


GENERAL CONCLUSIONS. 


As a result of the examinations made, second premolars and third 
molars cannot be regarded as providing adequate material for deter- 
mination, and no species founded on a second premolar or a third 
molar can be regarded as valid. If second premolars and third molars 
are known in a species from certain association with determinable 
teeth, corresponding teeth showing close agreement can then be 
assigned to the species with fair certainty. On general grounds of 
size and complexity, isolated second premolars and third molars may 
be referred to a species as “‘cf. species”’. 

In the remarks which follow, only the third and fourth premolars 
and the first and second molars are regarded as exhibiting characters 
of value in specific determination. 

The position of the tooth in the series affects two main characters 
as follows :— 


1. The transverse diameter of the crown, measured over the enamel, 
is about 5 per cent. greater in premolar than in molar teeth. 

2. The enamel pattern is somewhat more complex in the premolar 
than in the molar teeth, the fourth premolar showing the most complex 
and the first molar the simplest enamel folding. The mesostyle is 
somewhat reduced in size in the upper molars, and the entoflexid lobe 
is smaller in the lower molars than in the premolars. 


Bearing in mind the position of the tooth in the series, and the 
effect upon it of the above factors, the following generalisations 
may be made with regard to the definition and identification of 
equine species :— 


1. The transverse diameter (“‘breadth’’) measured across the enamel 
is a character of considerable value in determination. The normal 
departure in size is not more than 10 per cent. and the maximum 
variation 20 per cent. If the transverse diameter of a tooth departs 
by more than 20 per cent. from the measurement of the type, wt cannot be 
regarded as belonging to the same species. If several teeth of a species 
are known, the possible range which would exclude a specimen will 
be further reduced and a departure of 15 per cent. may be regarded 
with suspicion. It must be remembered, however, that different 
species overlap in point of size, and that size alone is not a sufficient 
basis for determination. 


422 Annals of the South African Museum. 


A. In Upper Cheek Teeth. 


The following factors may be regarded as of reasonable constancy 
and value in determination :— 


1. The relative anterior and posterior development and form of the 
protocone. (The absolute size of the protocone is subject to a variation 
between premolars and molars of as much as 20 per cent.) 

2. The form of the ectoloph and styles. 

3. The presence or absence of the pli-protoconule and pli-postfossette 
and their degree of development. 


The following factors may be regarded as of confirmatory value :— 


(a) The partial isolation of the hypocone (or the lack of such 
isolation). 

(b) The protocone ratio (subject to a variation of 30 per cent.). 

(c) The presence of any or all of the following folds, though their 
absence cannot be regarded as significant :— 


(i) Pli-hypostyle. 
(ii) Pli-caballin (particularly in true molars). 
(iii) Pli-protoloph (least reliable). 


(d) The form of the inner wall of the protoconule. 
(ec) The depth of the hypoglyph. 


B. In Lower Cheek Teeth. 


The following factors may be regarded as of reasonable constancy 
and value in determination :-— 


1. The shape of the outer walls of the protoconid and hypoconid. 
2. The relative development, degree and mode of separation of the 
metastylid and metaconid. 


The other factors which, though variable, may be regarded as of 
confirmatory value are: 


(a) The size and shape of the entaconid. 

(6) The relative sizes of the stylids and conids (if unusual). 

(c) The development of the ectostylid ridge or fold in the anterior 
wall of the hypoconid. 

(d) The size and form of the metaflexid and entoflexid lobes. 


A Critical Revision of the Quaternary Perissodactyla. 423 


Famity EQUIDAE. 


Genus EuRYG¢NaTHOHIPPUS van Hoepen 1930. 
Genotype: Hurygnathohippus cornelianus van Hoepen. 


Broad mandibular symphysis with the four first and second incisor 
teeth large, anteriorly flattened and arranged almost in a straight line, 
each incisor showing the cup or ‘‘Mark’’; the third incisors small and 
lying behind and in contact with the second incisor. (The cheek 
teeth are unknown.) 


Eurygnathohippus cornelianus van Hoepen. 


Eurygnathohippus cornelianus van Hoepen 1930. Pal. Nav. Nas. 
Mus. Bloemfontein, IT, 2, pp. 23-24, figs. 20-22. 


Type: Anterior portion of lower jaw with incisor teeth. Nas. Mus., 
No. C.679. (Fig. 8.) | 

Locality: Uitsoek, near Cornelia, Orange Free State. 

Horizon: The “Cornelia” Beds of van Hoepen (19304A). 


The following is a translation of van Hoepen’s description :— 


“There is preserved the anterior portion of the lower jaw. The four 
incisors are long and broad. The teeth lie deep in the antero-central 
part of the jaw and their grinding surface is practically horizontal. 
The teeth are anteriorly flattened. The first incisor has an indistinct 
groove along the centre of its anterior or lower surface. The second 
incisor has two such grooves. On the inner surface each tooth has 
two surfaces, ribbed parallel to the height, which meet in a blunt 
corner and a thick ridge. The canine is approximately one-third 
the width of the other teeth; unfortunately both are broken off. The 
four big incisors each show a large cup or mark, entirely surrounded 
by enamel and also completely filled by cement. The enamel on the 
front of the tooth is thick, but on the back it is thin. The two small 
canines show no mark; they are broken off very low down.” 


The present writer conjectured on the possibility of the teeth, which 
van Hoepen regarded as canines, being in point of fact reduced third 
incisors, and this suggestion was independently put forward by Dr. 
L. H. Wells to the writer. The writer and Dr. Wells have had the 
opportunity, through the courtesy of Dr. van Hoepen, of examining 
the type specimen, and are of the opinion that this is probably the 
case and that the specimen is that of a female individual, consequently 
lacking canine teeth. This reduction of the third incisors is not quite 
such a startling supposition as would be their complete absence, 


424 Annals of the South African Museum. 


though, as Dr. van Hoepen has said, it is but a step in that direction. 
The jaw thus clearly belongs to a horse, presumably a very large one, 


Fie. 8.—Upper, right lateral and lower views of anterior portion of 
mandible of type of Hurygnathohippus cornelianus van Hoepen. Two- 
thirds natural size. (From van Hoepen.) 


but there is some difficulty in absorbing it into Equus. Van Hoepen’s 
genus must stand in the mean time, and if “third incisor” be sub- 
stituted for “canine”’, his description cannot be bettered. 


A Critical Revision of the Quaternary Perissodactyla. 425 


Genus Notouipparion Haughton 1932. 
Genotype: Notohipparion namaquense Haughton. 


Rather low-crowned heavily cemented hypsodont lower cheek 
teeth with an extra antero-external cingulum fold or column, either 
isolated or fused with the parastylid,* present in all the permanent 
cheek teeth except the second premolar, and a deep groove separating 


the strongly developed metaconid and metastylid. The upper 
dentition is unknown. 


Notohipparion namaquense Haughton. 


Notohipparion namaquense Haughton 1932. Ann. 8. Afr. Mus., 
XXvll, pp. 421-423, fig. 5. 
Cotypes: Series lower Pm,, Pm,, Pm,, M, and M, of the left side 
and lower M, ofthe right side. S.A. Mus., No. 9982. (Fig. 9.) 
Locality: “40 miles east of Springbok in Namaqualand... .” 
Honzon: “from a granite gravel 60 feet down in the surface 
limestone”’. 


Measurements : 
LPm, LPm, LPm, LM, RM, LM, 
Breadth . ; 16 mm. 17mm. 17mm. 16mm. 14:5 mm. 12-5 mm. 
Length . - 295mm. 27mm. 26mm. 23mm. 24mm. 29mm. 
Height . - 145mm. 175mm. 25mm. 23mm. 30mm. 34mm. 


The following is Haughton’s description of the material :— 


“The lower teeth upon which this new genus is founded, indicate 
a stage of equine evolution not hitherto discovered in South Africa. 
They consist of a series from pm? to m!, together with m? of the left 
side, and pm‘, m* and m? and a part of m® of the right side, obviously 
of the same individual. As preserved the teeth are all low, the height 
not being much greater than the length. The main features can be 
distinguished from the drawings given. In all the teeth the meta- 
stylid column is separated from the metaconid column to the base of 
the crown, and the former projects further inwards than the metaconid 
or the entaconid, whilst its posterior flange overlaps the anterior 
border of the entaconid. The antero-external cingulum fold [proto- 
conid fold] is prominent to the grinding surface, being fused with the 
parastylid in pm? and m!, but still separated from it in pm‘, m? and 
m®, In pm‘ of the right side (but not of the left), in m' and m?, there 


* As defined by Osborn. Stirton (1941) replaces this by “‘paralophid”’ and uses 
‘“‘narastylid”’ for the antero-external column. Such reapplication of an existing 
name is too confusing to be accepted. Dr. L. H. Wells has suggested to the writer 
that this extra fold might be termed the “‘ protoconid fold”’. 


VOL. XXXI, PART 4. 36 


426 Annals of the South African Museum. 


is an ectostylid pillar, which near to the root fuses with the wall of 
the hypolophid to form an ectostylid ridge. Enamel fairly wavy. 
Teeth heavily cemented” (pp. 421, 422). 

“Unfortunately nothing is known of the sequence of gravels and 
limestone encountered in the well from which Notohipparion nama- 
quense was obtained, and but little hght can be thrown upon its age. 
The valleys of Namaqualand seem to have suffered a progressive 
infilling with sand from Upper Cretaceous times onward; but the 
process was, in all probability, not a continuous one and further study 
will probably reveal breaks in the sedimentation. In so far as com- 
parison is possible, Notohipparion would seem to represent an early 
Pliocene stage of equine evolution” {p. 425). 


In view of the fact that teeth of another member of the Hipparion 
group occur in the Vaal River gravels associated with well-made stone 
implements, it would appear that these forms have survived in this 
region until a much later time than elsewhere. Though Notohipparion 
is quite possibly a Pliocene form, it need not necessarily be “early 
Pliocene” as Haughton suggests, and it is included here since its 
horizon is so uncertain and since it may have a bearing on the 
undoubted Pleistocene forms. 


Genus STYLOHIPPARION van Hoepen 1932. 
Genotype: Stylohipparion steytleri (van Hoepen). 
(=8S. hipkini van Hoepen). 


High-crowned rather narrow hypsodont lower cheek teeth with a 
strongly developed isolated pillar external to the ectosylid, possessing 
no external groove between hypoconid and hypoconulid, having 
narrow protoconid and hypoconid and small rather widely separated 
metaconid and metastylid. 

High-crowned upper cheek teeth with isolated oval protocone and 
possessing a small flange on the antero-internal side of the parastyle. 


Stylohipparion steytlert (van Hoepen). 


Hipparion steytlerr van Hoepen 1930. Pal. Nav. Nas. Mus. Bloem- 
fontein, II, 2, pp. 21-23, figs. 14-19. 

Stylohipparion hipkint van Hoepen 1932. Pal. Nav. Nas. Mus. 
Bloemfontein, IT, 3, pp. 31-32, figs. 14-17 and 18-20. 

Stylohupparion steyilery van Hoepen 1932. Ibid., pp. 33-35, figs. 
21-23. 


A Critical Revision of the Quaternary Perissodactyla. 427 


Fig. 9.—Series*of lower cheek teeth of the type of Notohipparion 
namaquense Haughton. Natural size. (From Haughton.) 


428 Annals of the South African Museum. 


Type: Upper M! or M? of the right side. Nas. Mus., No. C.558. 
(Fig. 10.) 

Locality: Uitsoek, near Cornelia, O.F:S. 

Horizon: The “‘Cornelia Deposits”’ of van Hoepen. 


Measurements: 
Breadth . ; « 22 7am, 
Length . ; . 22mm. 
Height . . . 54mm. 


Paratypes: Upper M3 of the left side. Nas. Mus., No. ©.555. 
(Fig. 10.) 


Lower M, and M, of the left side. Nas. Mus., No. 
C.556. (Fig. 10.) 


Measurements: 
LM LM, LM, 
Breadth . . 18mm. 10(12) mm. 10-5(12-5) mm. 
Length . . 21 mm. broken 21 mm. 
Height . . 66 mm. 32 mm. 41 mm. 


(The figures in brackets include the accessory outer column.) 


Upper Teeth. 


The halves of the ectoloph are concave inwards though the posterior 
half may be flattened. The metastyle is small and the mesostyle and 
parastyle narrow, the latter having an unusual anterior flange or 
groove. The protocone is isolated and oval in shape with a somewhat 
flattened interior face. A small tongue projects into the medivallum 
towards the protocone. The hypocone is very small, being less than 
one-third the size of the protocone. A deep, sometimes double 
protoconule fold is present and, with the prefossette folds, tends to 
isolate the postero-internal corner of the prefossette. Protoloph, 
hypostyle and postfossette folds are present and are of moderate depth. 


Secondary plications are present in the region of the prefossette fold 
and appear to be persistent. 


Lower Teeth. 


Protoconid and hypoconid are rather narrow transversely and have 
slightly flattened or even concave outer walls. An ectostylid ridge 
or fold is present in the anterior wall of the hypoconid, and external 
to this is a stout oval pillar arising from the cingulum and reaching 


A Critical Revision of the Quaternary Perissodactyla. 429 


Fie. 10.—Stylohipparion steytlert (van Hoepen). 
Top: Type upper right second (or third) molar; crown and anterior views. 
Left: Paratype upper left third molar; crown and outer lateral views. 
Right: Paratype lower left first and second molars; inner and crown views. 
All natural size. (From van Hoepen.) 


almost to the unworn crown. There is no external groove marking off 
hypoconid and hypoconulid. The entaconid is small, and the meta- 
conid and metastylid are rather small and widely separated by a 
rounded groove. The entoflexid lobe is only slightly longer than the 


430 Annals of the South African Museum. 


metaflexid lobe, and folding in their walls is slight or lacking, a small 
notch in the anterior of the entoflexid lobe being sometimes present. 


Referred Upper Teeth. 

An upper second (or possibly third) molar of the left side has been 
recovered from the Vaal River deposits at Christiana, probably from 
Younger Gravels. This specimen (Arch. Sur., No. 113) is consider- 
ably worn, but retains all the characters shown by the less worn type 
and paratype. 


Referred Lower Teeth. 


Five lower teeth, being a series from LPm, to LM, from the type 
locality, were referred to this species by van Hoepen (1932), and the 
enamel pattern of four of them is shown in fig. 11. 


bat) 

\ 

(ina Ue a 
E BA OOOCERM 


Fic. 11.—Referred series of lower left cheek teeth of Stylohipparion steytlert 
(van Hoepen). Crown views. Natural size. (From van Hoepen.) 


The following are the dimensions of these teeth (as preserved) :— 


LPm, LPm, LM, LM, LM, 
Breadth . Samim: 12 (14) mm. 12 mm. 11-5 mm. 9 mm. 
Length ; . 25mm. 28 mm. 22 mm. 24-5 mm. 23 mm. 
Height ; . 64mm. 74 mm. 69 mm. 73 mm. 38 mm. 


A lower second molar from the type locality was described by van 
Hoepen in the same paper (1932) as the type of a new species, 
S. hipkin. The specimen (Nas. Mus., No. 0.797) is very little worn 
and thus exhibits an entirely uncharacteristic pattern. Its height 
is 81 mm., and 15 mm. below the exposed grinding surface the 
dimensions of the tooth are: breadth 12-5 mm., length 26-5 mm.; 
in the middle of the tooth the corresponding dimensions are 13 mm. 
and 22mm. A left second premolar is provisionally referred to this 
new species, but there does not appear to exist any valid reason for 
separating either of these teeth from the earlier species. 

Two lower teeth recovered by Broom from the Kromdraai Cave 
have been referred by the present writer to this species but they have 
not yet been described. 


A Critical Revision of the Quaternary Perissodactyla. 431 


Genus Equus Linnaeus 1758. 
Genotype: Equus caballus* Linnaeus. 
Equus burchellii* (Gray). 


Asinus burchelliz* Gray 1824. Zool. Journ., 1, p. 247, peo, aigs, 1,2. 

Equus quagga wahlbergi* of most authors. 

Equus platyconus van Hoepen 1930. Pal. Nav. Nas. Mus. Bloem- 
fontein, IT, 1, pp. 4-5, figs. 3-5. 

Equus simplicossimus van Hoepen 1930. Ibid., p. 6, fig. 7. 

Equus simplicissimus van Hoepen 1930. Pal. Nav. Nas. Mus. 
Bloemfontein, II, 2, p. 21, figs. 12, 13. 

Kraterohippus elongatus van Hoepen 1930. Pal. Nav. Nas. Mus. 
Bloemfontein, II, 1, pp. 7-8, fig. 9. 

Equus lyle: Dreyer 1931. ‘‘New Fossil Mammals and Man”, p. 30, 
poy, we. 9: pl. vi, fies. 6,7, 8; pl: vil, fig) 8. 

The general dental formula is the same as in Equus caballus*, but 
the vestigial first upper premolar, which is normally lacking in £. 
caballus* and in most other members of the genus, is frequently 
present in £. burchellw* and is shed only comparatively late in life. 
The upper incisor teeth closely resemble those of H. caballus*, but 
the lower incisors all typically lack the cup or mark so characteristic 
of the commor horse. This cup is normally formed as a result of 
closure of two posterior folds of the tooth, and its incomplete formation 
is well seen in many specimens of LH. asinus*. In E. burchellii* it is 
completely absent in the lower I,, partially formed in I, and partially 
or completely formed in I,, but even if present in I, is so shallow as 
rapidly to disappear with wear. A typical specimen is shown in fig. 12. 
This feature is not characteristic of either H. zebra* or E. quagga*, 
though in the latter species the cup is often absent in the lower third 
incisor, and is not as deep as in EH. caballus*. The retention of the 
vestigial first premolar and the nature of the incisor teeth may both 
be regarded as rather primitive characteristics. 

The upper cheek teeth are a good deal smaller than those of HL. 
caballus*, and the dimensions of a typical complete dentition (Tvl. 
Mus., No. 173, fig. 13c) in normal wear are: 


Pm? Pm? eem* M! M2 M$’ 
Breadth . 22mm. 24mm. 24:5mm. 23mm. 23mm. 21 mm. 
Length ~. 85mm. 24mm. 25mm. 205mm. 2mm. 23mm. 


The breadth variation noted in Pm*-Pm* is 22-26 mm. and in 
M2-M? is 21-25 mm. 


432 Annals of the South African Museum. 


Fic. 12.—Lower incisor teeth of Equus burchellii* (Gray) to show the 
characteristic absence or reduction of the cup or “mark”. Natural 
size. (Original.) 


A Critical Revision of the Quaternary Perissodactyla. 433 


The halves of the ectoloph are concave from without inwards and 
curve smoothly into the styles, though the mesostyle may overhang 
slightly. The parastyle is commonly obliquely flattened anteriorly 
in the premolars but less commonly so in the true molars. This and 
the mesostyle are sometimes grooved externally for a small extent 
below the grinding surface. The metastyle is small and not prominent. 
The protocone is elongate oval in form (proportionally narrower than 
in H. caballus*), and the portion anterior to the junction with the 
‘protoconule comprises about one-third of the total protocone length 
(except in Pm? where the anterior extent is very small in all species of 
Equus). The hypocone is not large and frequently shows a slight 
bulge into the medivallum, giving to it the appearance of a tendency 
towards isolation from the metaconule. The caballine fold is typically 
absent, but may be present in some individuals, particularly in the 
premolars. The fossette folds are generally small except for a well- 
developed pli-protoconule and a fairly good pli-postfossette. The 
pli-hypostyle and pli-prefossette are very variable in development, 
but the pli-protoloph is usually present and often fairly deep. The 
crown patterns of three typical dentitions are shown in fig. 138. 

The lower teeth are also a good deal smaller than those of E. 
caballus*. The dimensions of the lower teeth (fig. 14) in the individual 
for which the measurements of the upper teeth have already been 
given are: 


Breadth f ~ 125mm. 13-5mm. 14mm. 12mm. 11:-5mm. 11 mm. 
Length ; 2 228mm. 25mm. 24mm. 22mm. 2mm. 27 mm. 


The breadth variation noted in Pm,-Pm, is 12-15 mm. and in 
M,—M, is 11-14 mm. 

The outer walls of the protoconid and hypoconid are both flattened 
or even concave inwards, and the ectostylid fold in the hypoconid is 
small or even absent. The entaconid is small and somewhat quadrate 
inform. The metaconid is oval, and is separated from the metastylid 
by a fairly sharp-pointed groove. The metastylid itself is rather 
small, pear-shaped and bluntly pointed. The metaflexid lobe is 
smaller than that of the entoflexid, and the outer walls of the ento- 
flexid may show some folding which does not persist with advanced 
wear. 


Referred Fossil Material. 


Petrified specimens of this species are of common occurrence In cave 
deposits, in superficial deposits, in river gravels and elsewhere. Such 


A Critical Revision of the Quaternary Perissodactyla. 435 


@ 


Fie. 14.—Right lower cheek teeth of Hquus burchellii* (Gray). The 
lower dentition C belongs to the individual figured in fig. 13, C. Natural 
size. (A, after Owen; B and C original.) 


436 : Annals of the South African Museum. 


material does not always seem to be of very great antiquity and is 
quite possibly only of Upper Pleistocene Age. 

The species Equus platyconus van Hoepen and Kraterohippus 
elongatus van Hoepen have already been regarded by Haughton 
(1932) in his revision as probable synonyms of the living Burchell’s 
zebra, and there seems no reason to doubt the correctness of this 
conclusion. Kraterohippus elongatus, moreover, is founded on a 
broken upper Pm?, almost certainly incorrectly restored and is value- 
less as a type. Equus sumplicissimus van Hoepen also appears to 
fall well within the range of variation of the living species and does not 
warrant distinction. 

Equus lylet Dreyer, founded on well-mineralised material from the 
springs at Floris Bad, was regarded by Haughton as falling within 
the range of variation of the living species. This is certainly the case, 
most of the specimens from the type and neighbouring areas, however, 
lying close to the lower size limit of the species. Their enamel 
characters are quite typical and the material does not apparently 
require specific distinction, though a varietal or sub-specific separation 
may prove to be justified. 

Mineralised specimens of Equus burchelliv* occur in the deposits of 
the Vaal River basin. 


Equus zebra* Linnaeus. 


Equus zebra* Linnaeus 1758. Syst. Nat., Ed. 19, 1, 


The incisor teeth have been considered under LZ. burchellir* above. 

The upper cheek teeth differ very little in size from those of E. 
burchellu*, though on the average they are possibly a millimetre 
broader than the average bontequagga. The halves of the ectoloph 
are almost straight or even slightly convex, and the parastyle and 
mesostyle are somewhat abruptly marked off from the ectoloph walls. 
These features are less apparent in the true molars than in the pre- 
molars. The protocone is sub-triangular rather than oval, and the 
portion anterior to its junction with the protoconule is less than one- 
third of the protocone length. The parastyle is commonly flattened 
anteriorly but is not as obliquely directed as in #. burchellu*. The 
hypocone is only a little smaller than the protocone. A caballine 
fold is often, but not consistently, present. The fossette folds are 
all small, and even the pli-protoconule and pli-postfossette may be 
virtually lacking. The pli-protoloph is typically absent or extremely 
small. 


438 Annals of the South African Museum. 


The lower teeth are also comparable in size with those of the 
bontequagga. The outer walls of the protoconid and hypoconid are 
convex and well rounded, and the ectostylid fold is small or lacking. 
The entaconid is rounded, as also is the metaconid, but the metastylid 
is pear-shaped and rather pointed. The groove separating metaconid 
and metastylid is somewhat pointed. The metaflexid and entoflexid 
lobes are simple. 

Two upper and a lower dentition are shown in fig. 15. 


Referred Fossil Material. 


The only petrified specimens certainly to be ascribed to this species 
were recovered from the Cango Caves near Oudtshoorn, but the 
remains of zebrine skulls are not uncommon in the older dune sands 
of the coastal belt in the Cape Province. 


Equus quagga* Gmelin. 


Equus quagga* Gmelin (Linnaeus 1788). Syst. Nat., Ed. 13, 1, p. 21. 

Equus quagga* Cooke 1941. S. Afr. J. Sci., xxxvui, p. 307. 

Equus quagga* Shapiro 1943. 8S. Afr. J. Sci., xxxix, p. 117. 

(non E. quagga quagga* Dreyer 1931. “New Fossil Mammals and 
Man”’, p. 33.) 

(non EF. quagga* var. Haughton 1932. Ann. 8. Afr. Mus., xxviii, 
p. 424.) 


Equus quagga* became extinct in 1872 before most museums had 
commenced the collection of skeletal material from Southern Africa, 
and only a cast in the Transvaal Museum, taken from a skull in the 
possession of the Stuttgart Museum, represents a specimen actually 
known to be that of the true quagga. This individual was very 
young, and the cast is not of great value for identification purposes. 
Fortunately the great Richard Owen, in the “Philosophical Trans- 
actions of the Royal Society of London” for 1869, gives natural size 
figures of the dentition of a good adult specimen at that time in the 
Royal College of Surgeons. Dr. Robert Broom, while in London 
many years ago, made drawings of the teeth of a quagga in the British 
Museum collections, and these agree with those given by Owen 
in all characters. Some old skulls and jaws in the McGregor 
Museum, Kimberley, and in the collections of the Kingwilliamstown 
Museum, agree closely with the figures published by Owen and with 


A Critical Revision of the Quaternary Perissodactyla. 439 


the manuscript drawings given to the writer by Dr. Broom, and as 
these skulls were collected in a region certainly at one time overrun 
by quagga, this material may with fair certainty be taken to represent 
the species. Furthermore, the characters which Owen remarks as 
distinguishing the quagga from the other two South African species, 
are present in these skulls. 

From the limited material available, it would appear that the teeth 
of this species do not differ very markedly in size from those of the 
zebra and bontequagga. The largest and smallest upper dentitions in 
the McGregor Museum collection (fig. 16, B, C) have the following 
dimensions over the enamel :— 


Pm? me Pm* WE M? M3 
Breadth 215mm. 225mm. 22mm. 205mm. 20:0mm. 19-5 mm. 
Length 305mm. 235mm. 215mm. 20:(0mm. 195mm. 20:5 mm. 
Breadth 23-0 mm. 270mm. 265mm. 245mm. 240mm. 19-5 mm. 
Length 325mm. 245mm. 245mm. 215mm. 215mm. 23-5 mm. 


Owen’s figure gives measurements only slightly smailer than the 
dimensions of the larger dentition, so that it would appear that the 
average quagga dentition was a little larger than the normal bonte- 
quagga, and the range in breadth is somewhat greater as well, being 
22-27 mm. for Pm?—-Pm*# and 20-25 mm. for M!-M?. 

The halves of the ectoloph are almost straight or even slightly 
convex, and are abruptly marked off from the parastyle and mesostyle 
by an acute angle, though this character is suppressed in the third 
molar and not clear in the second premolar. The external face of the 
parastyle, unlike that in EL. burchelli* and EL. zebra*, is not directed 
in an oblique angle anteriorly but lies roughly parallel to the axis of 
the row of cheek teeth. The protocone is elongate oval or somewhat 
triangular in form, and commonly has an almost median indentation 
in its inner wall, giving a rather bilobate appearance. The junction 
of the protocone with the protoconule is only slightly anterior to the 
middle of the protocone. The hypocone is small and the hypoglyph 
is rather variable. The caballine fold is normally absent. The 
fossette folds are all small, even the pli-protoconule and pli-postfossette 
being commonly reduced to a small notch only. Nevertheless the 
pli-protoloph and pli-hypostyle are both normally visible, though 
the pli-prefossette is typically small or absent. The crown patterns 
of three upper dentitions are shown in fig. 16. 

The lower teeth are comparable in size with those of HL. burchelli*, 
two typical dentitions measuring: 


A Critical Revision of the Quaternary Perissodactyla. 441 

Pm, Pm: Pm, M, M, M, 
Breadth 130mm. 140mm. 145mm. 125mm. 12:0mm. 11-5 mm. 
Length 275mm. 25-0mm.. 240mm. 225mm. 220mm. 26-0 mm. 
Breadth 130mm. 140mm. 140mm. 125mm. 12-0 mm. 11-0 mm. 
Length 315mm. 265mm. 260mm. 235mm. 240mm. 23-5 mm. 


Owen’s figure agrees very closely in dimensions with those of the 
latter species. The probable range in breadth may be estimated as 
12-16 mm. for Pm,-Pm, and 11-15 mm. for M,-M,,. 

In character the lower teeth resemble those of bontequagga 
rather than zebra, for the outer walls of the protoconid and hypoconid 
are somewhat flattened, the latter more so than the former. The 
ectostylid fold is often present. The metaconid is rounded or oval, 
and is separated from a pear-shaped metastylid by a rounded groove. 
The entaconid is rounded to quadrate and notably larger in the 
premolars than in the molars. 

Three dentitions are shown in fig. 17. 

Probably largely in consequence of the confused ideas which have 
hitherto existed regarding the characters of the teeth of Equus 
quagga*, none of the teeth referred to by various authors as “Z. 
quagga”’ or “ E. quagga quagga”’ can be regarded as actually belonging 
to Gmelin’s species. These so-called “quagga” teeth are mainly 
those of £. burchellia* (Gray) but, remarkably enough, none of the 
fossil teeth described under other names can be regarded as belonging 
to E. quagga* either, so that it would appear that this recently extinct 
species has not hitherto been recognised to any notable degree in our 
fossil collections. This species is represented by material in the 
McGregor Museum, Kimberley, derived from surface deposits at 
Koffiefontein, Orange Free State. It has also been recorded from a 
cave near Kuruman (Cooke, 1941), from Bankies, O.F.S. (Shapiro, 
1943) and appears to be present in the deposits of the Vaal River valley. 
Doubtless more material will be recognised in due course. 


Equus capensis Broom. 


Equus capensis Broom 1909. Ann. 8. Afr. Mus., vil, pp. 281-282. 

Equus capensis Broom 1913. Bull. Amer. Mus. Nat. Hist., xxxii, 
p. 437, fig. 1. 

Equus capensis Broom 1928. Ann.S8. Afr. Mus., xxii, p. 441, fig. 2, A. 

Equus cawoodi Broom 1928. Ibid., pp. 443-444, fig. 3, A. 

Equus gigas van Hoepen 1930. Pal. Nav. Nas. Mus. Bloemfontein, IT, 
1, pp. 2-3, fig. 1. 

Equus capensis Dreyer 1931. Ibid., pp. 36-87, pl. vii, fig. 5. 

VOR, XXXI, PART 4. 37 


442 Annals of the South African Museum. 


< a = 


Fic. 17.—Right lower cheek teeth of Equus quagga* (Gmelin). Natural size. 
(A, after Owen; B and C original.) 


A Critical Revision of the Quaternary Perissodactyla. 443 


Equus westphali Dreyer 1931. Ibid., pp. 36-37, text-fig. 

Equus capensis Haughton 1932. Ann. 8. Afr. Mus., xxviii, pp. 410- 
412, fig. 2. 

Equus capensis Cooke 1939. 8. Afr. J. Sci., xxxvi, pp. 413-414, 
fig. 2. 

Equus capensis Cooke 1941. 8. Afr. J. Sci., xxxvii, pp. 308-311, 
“figs. 4, 5. 

Equus capensis Wells and Cooke 1942. Trans. Roy. Soc. S. Afr., 
XX1X, pp. 228-229, fig. 12. 

Equus capensis Shapiro 1943. 8. Afr. J. Sci., xxxix, pp. 117-178. 

(non #.cawoodi van Hoepen 1930. Pal. Nav. Nas. Mus. Bloemfontein, 
II, 1, pp. 3-4, fig. 2.) 

(non £. cawoodi Dreyer 1931. “New Fossil Mammals and Man”’, 
pp. 26-29, pl. vi, figs. 8-12; pl. v, fig. 2.) 

(non £. capensis Broom and Le Riche 1937. S. Afr. J. Sci., xxxiil, 
pp- 769-770, fig. 1.) 

(non pars H. capensis Cooke 1939. S. Afr. J. Sci., xxxvi, pp. 413- 
414, fig. 2a.) 

Type: Series lower left P,, P;, P,, M,, M, embedded in limestone 
and damaged on the inner sides. 8.A. Mus., No. 658. (Fig. 18.) 
Locality and Horizon; Beach debris, Yzerplaats, Maitland, Cape 
Province. 


Measurements: 
ibles LP, GP; LM, LM, 
Breadth (damaged) 2 215+ mm. 15+ mm. 14+ mm. 14+ mm. 
Breadth (as 
restored) . : 2 19-5 mm. 18-5-19 mm. 18mm. 17-5 mm. 
Length : . ?30 mm. ?35 mm. 34:5 mm. 31mm. 30mm. 
Height : a? (Og. 87 mm. 103 mm. 92mm. 94mm. 


Neotype (Haughton 1932): Lower left fourth premolar. §S.A. Mus., 
No. 2821. (Fig. 18.) 
Locality: Saldanha Bay. 


Measurements: 
LPm, 
Breadth Z é . 18-5 mm. 
Length : A - 33-0 mm. 
Height : ‘ . 92-0 mm. 


The type of this species was first described by Broom in 1909 but 
was not figured until 1928. It comprises a damaged series of teeth 
embedded in a slab of sandy limestone found on the beach at Yzer- 
plaats, Maitland district, Cape Province. It would appear that this 
limestone represents a former land deposit now lying below sea-level 


444 


Annals of the South African Museum. 


Fic. 18.—Equus capensis Broom. 
Crown view of the type series of left lower cheek teeth as embedded in 


Top: 
the limestone matrix. (Original.) 
Centre: Restoration of crown patterns of type series. (Original.) 
Below: Enamel pattern of neotype lower left fourth premolar. (From Haughton.) 


All natural size. 


ae 


A Critical Revision of the Quaternary Perissodactyla. 445 


as a result of late crustal warping. While it is impossible to arrive 
directly at an estimate of the age of the original deposit, this must 
antedate the warping which itself is most probably the final crustal 
warping occurring between the development of the “ major emergence”’ 
and the “minor emergence” raised beaches. The dating of this event 


is difficult, but the writer has elsewhere (Cooke, 1941 ) suggested that 


it coincides approximately with the existence of the Stellenbosch 
industry and may be broadly Middle Pleistocene. 

- If this estimate is not far wrong, the Equus capensis bearing lime- 
stone belongs most probably to the Middle Pleistocene, and the general 
affinities of the teeth also support a Middle Pleistocene age. 

The teeth themselves are exposed on a fractured surface of the slab, 
and the inner portions of each tooth have been lost. The series 
belongs to the left lower jaw and lacks the third molar, but the second 
premolar is completely embedded and the anterior portion of the third 
premolar partly covered by limestone. The patterns of the grinding 
surfaces are consequently visible only in part of the third premolar, 
the fourth premolar and the first and second molars, but all the teeth 
are well exposed in their inner lateral aspects. Broom figured a 
restoration of these four teeth in 1928, but many features of the 
drawing are faulty, and the restored portions probably incorrectly 
given. Fig. 18 shows an accurate drawing of the enamel patterns as 
they appeared in the slab in 1940, though possibly the specimen is now 
less perfect than in 1928. Nevertheless, Broom’s restoration fails to 
show the distinct flattening of the hypoconid outer wall, gives in- 
correctly the shape of the visible portions of the entaconid and 
simplifies the irregularities in the entoflexid. As Haughton has 
pointed out: “In his restoration of the missing portions he has 
shown the metaconid, metastylid and entoconid as being rounded in 
outline, his restoration of the metastylid in particular differing con- 
siderably from the somewhat triangular or pointed form seen in 
Equus caballus.’’ It is impossible from the specimen to see any 
grounds for maintaining this restoration, and in the light of all the 
available evidence a new restoration is now given which is probably 
not far from the truth (fig. 18). 

In form these teeth resemble those of the true quagga rather than 
the other living species, but they are notable for their very large size, 
far exceeding the dimensions of any living type. The protoconid and 
hypoconid are well developed, and the outer wall of the latter is more 
markedly flattened than in the former. The protoconid projects, in 
the molars at least, beyond the outer wall of the hypoconid, giving a 


446 Annals of the South African Museum. 


somewhat lopsided appearance to many of the teeth. The ectostylid 
fold in the hypoconid wall is small or completely lacking. The 
hypolophid is quite distinctly marked off from the hypoconid. The 
entaconid is quadrate to semi-lunate in form with a distinctly angular 
antero-internal corner (“entostylid”). The metastylid is pear- 
shaped and somewhat pointed (if the restoration or evidence from the 
neotype and other specimens can be assumed to be correct). The 
metaconid is oval and is separated from the metastylid by a pointed 
groove. The outer wall of the entoflexid is irregular or wavy in the 
premolars and slightly concave in the molars. The simplicity of the 
molars as compared with the premolars, and also the rather great 
disparity of form and size, is striking in some of the series available. 
The range in breadth for the premolars appears to be about 18-5 to 
22 mm. and for the molars 17-5 to 21 mm. 

Amongst the large number of lower teeth referred to this species, 
the best preserved and most valuable series is shown in Fig. 19. 
These teeth come from a thermal spring at Vlakkraal near Bloem- 
fontein, Orange Free State, and have been described by Wells and 
Cooke (1942). Their individual dimensions are as follows:— 


Pm, Pm, Pm, M, M, 
Breadth . 5 : - 185mm. 19mm. 195mm. 17mm. 17-5 mm. 
Length . : ; : 38mm. 34 mm. 34mm. 29mm. 30-5 mm. 
Height (excluding roots) . 56mm. 62 mm. 74mm. 58mm. 51+ mm. 


The teeth are thus very slightly smaller than those of the type series, 
but there can be little doubt regarding their close agreement in all 
essentials. This series also illustrates very well the lateral aspects of 
the teeth and their normal angles of wear, and is very useful as an 
aid in determining the position of an individual tooth in a series. 

The upper teeth of Equus capensis are so far not certainly known, 
but from the normal relationships between upper and lower teeth the 
expected breadth of the uppers would be 29 to 35 mm. for the pre- 
molars and 27 to 33 mm. for molars. Broom (1913 6, 1937), Dreyer 
(1931) and Haughton (1932) have referred large upper teeth to £. 
capensis. New species within the correct size group have also been 
erected by Broom (1928), van Hoepen (1930), Dreyer (1931) and Wells 
(1941). All these finds have been on open sites where association is 
virtually valueless, but material described by the writer (Cooke 
1941 a) and by Wells and the writer (1942) from sealed sites may 
elucidate the problem. 

In a large collection of equine remains from the Wonderwerk Cave 


is . 
cee 

. 5 Oyo of as 

ee ee 
ae ee 5 


“st; Ae? ere 


diiteutelats Tinga 


al “9.%°% 0 < 

CC tas ae) “g¢ =. 
oie Cer ae iy eee 

ee oS ce e@* .? ° 


ar 


Fig. 19.—Fine series of lower left cheek teeth referred to Equus capensis 
Broom. Outer lateral and crown views. Natural size. (From Wells, 
Cooke and Malan.) 


448 Annals of the South African Museum. 


in the Kuruman district, twelve teeth (upper and lower) were immedi- 
ately distinguished by their great size. Five are lower teeth which 
agree in pattern with those of H. capensis, though they are very 
slightly smaller than the type. The associated upper teeth can 
hardly belong to a different species, and these are closely comparable 
with the type of Equus cawoodi described by Broom (1928) from the 
Vaal River gravels. The type of EZ. cawoodi (which is most probably 
a Pm? and not a Pm‘ as Broom states) is little worn, and agree in 
almost every detail with a little worn LPm‘* from Wonderwerk. The 
Wonderwerk LPm¢ was sectioned, and on its sectional surface showed a 
very considerably simplified pattern agreeing very well with the uppers 
referred to EH. capensis by Broom (1913) and one referred to that 
species by Haughton (1932). Additional material from Wonderwerk 
described by Wells (1943) confirms this view. The large upper teeth 
associated with the fine lower series from Vlakkraal are very worn, 
but they agree in all essentials with EH. cawoodi and there can be little 
doubt that the two species are synonyms. Shapiro (1943) dissented 
from this view, but his argument was based chiefly upon the material 
erroneously referred to LH. capensis by Broom & le Riche and now falls 
away, as he himself would doubtless agree. 

The type RPm?® of HL. cawoodi (M.M.K., 3711) has the followine 
dimensions (fig. 20) :— 


Breadth, on grinding surface (which is Fi little 


worn) . 31-5 mm. 
Breadth, 3 cm. below grinding surface . : ; 32 mm. 
Length , ; : . : : : . 33 mm. 
Height ‘ é , : : : ; ‘ 80 mm. 


It may conveniently be taken as a neotype of the upper dentition 
of E. capensis. 

The form of the upper dentition of E. capensis (as now understood) 
is very similar to that of E. burchelli*, of which it is virtually an 
enlarged version. The halves of the ectoloph are concave inwards 
and curve easily into the styles, except for a distinct tendency for 
an anterior overhang of the mesostyle. The parastyle is obliquely 
flattened anteriorly in the premolars but less noticeably so in the true 
molars. The metastyle is small. The hypocone is moderately small 
but the hypoglyph may be rather deep. The caballine fold is 
sometimes present but may often be absent. The pli-protoconule 
and pli-postfossette are well marked, but the pli-protoloph and pli- 
hypostyle are very small and may disappear completely with wear. 
The pli-prefossette and secondary small postfossette are distinct in 


Upper left: 
Upper right: 
Lower left: 


Lower right: 


pau0ij9as 


Fie. 20.—Hquus capensis Broom. 

Crown and outer lateral views of neotype upper right third pre- 
molar. (Original.) 

Enamel pattern of two referred upper left fourth premolars from 
Saldanha Bay. (After Broom.) 

Enamel pattern of referred upper left second molar from Bloem- 
bosch, Darling. (After Haughton.) 

Original crown and sectioned enamel pattern of a broken, little 
worn, referred upper left fourth premolar from Wonderwerk. 
Sectioned surface approximately 2 cm. below crown. (From 


Cooke.) 
All natural size. 


450) Annals of the South African Museum. 


very early wear but vanish rapidly with abrasion. The protocone is 
elongate oval in form, with rather more than one-third of its total 
length lying anterior to the junction with the protoconule. The 
range in size is, as far as observation goes, that deduced from the 
dimensions of the lowers. 

Fig. 20 shows the form and pattern of the neotype (previously the 
type of EH. cawoodi), the two teeth referred to the species by Broom 
(1913), that referred to it by Haughton (1932), and the sectioned 
damaged tooth from Wonderwerk. Fig. 21 gives the pattern of part 
of a series from Wonderwerk, two teeth from Floris Bad and one from 
the Vaal River gravels. These figures give a good idea of the 
characters and variations of the species. 

The series of milk molars and two molars described by van Hoepen 
(1930 a) as Equus gigas do not appear to warrant specific distinction 
and may be referred to EL. capensis. The upper teeth from Vlakkraal 
agree with those from Floris Bad to which Dreyer gave the name 
E. helmet, a species which Haughton (1932) regarded as a synonym of 
E. cawoodi. This is accordingly a synonym of £. capensis, as also is 
E. westphali, which Dreyer erected on lower teeth of large size from 
the Vaal River gravels at Pniel. 

The tooth which van Hoepen (1930 a) referred to EL. cawoodi and 
subsequently (1930 b) to a new species Z. louwi does not appear to be 
E. capensis, but rather E.kuhni Broom. The same incorrect identifica- 
tion appears to apply to Dreyer’s FZ. cawoodi (1931) and to the specimen 
described and wrongly figured by Broom and le Riche (1937). The 
smaller of the two right lower fourth premolars referred by the present 
writer (Cooke 1939) to #. capensis seems in the light of later knowledge 
to belong to EZ. harrisi Broom. 


Equus kuhni Broom. 


Equus kuhni Broom 1928. Ann. 8. Afr. Mus., xxii, p. 444, fig. 3, B. 

Equus cawoodi van Hoepen 1930. Pal. Nav. Nas. Mus. Bloemfontein, 
II, 1, pp. 3-4, fig. 2. 

Equus louwi van Hoepen 1930. Pal. Nav. Nas. Mus. Bloemfontein, 
II, 2, pp. 19-21, figs. 6-11. 

Equus kuhni Haughton 1932. Ann. 8. Afr. Mus., xxviii, p. 414. 

Equus kuhni Cooke 1941. 8. Afr. J. Sci., xxxvii, pp. 307-308, fig. 3. 

Equus capensis Broom and Le Riche 1937. S. Afr. J. Sci., xxxiii, 
pp. 769-770, fig. 1, A. 


A Critical Revision of the Quaternary Perissodactyla. 451 


Fic. 21.—Hquus capensis Broom. 
Left: Part of incomplete series of referred left upper cheek teeth from 
Wonderwerk Cave. (From Cooke.) 
Referred left upper fourth premolar and second molar from Floris- 
bad. (After Dreyer.) 
Lower right: Referred upper right fourth premolar from Vaal River gravels at 
Austin’s Rush, Barkly West. (Original.) 
All natural size. 


Top right: 


452 Annals of the South African Museum. 


Sectioned 


Fig. 22.—Equus kuhni Broom. 
Left: Type upper right fourth premolar. (Original.) 
Right: Neotype upper left fourth premolar from Wonderwerk Cave, showing 
pattern on little worn crown and on surface sectioned 2-5 cm. below the 
crown. (From Cooke.) 


Natural size. 


Type: Upper Pm‘ of the right side. M.M.K., No. 3929. (Fig. 22.) 
Locality: Pniel, Vaal River. 7 
Horizon: % Vaal River gravels. 


Measurements: 
RPm#? 2 cm. below crown 
Breadth : : . w29:>)mm. 28-5 mm. 
Length : : . 431-0! 30-5 mm. 


Height : : + 273 -0imm.: 


A Critical Revision of the Quaternary Perissodactyla. 453 


Neotype: Upper Pm!‘ of the left side. Arch. Sur., No. 143. (Fig. 22.) 
Locality: Wonderwerk Cave, Kuruman District. 
Horizon: Disturbed cave deposits. 


Measurements: 
LPm#4 2-5 cm. below crown 
Breadth : 5 . 27-5 mm. 28 mm. 
Length é 5 - 29-5 mm. 28-5 mm. 
Height : 2 - 77-0 mm. 


The type of this species, like that of E. cawoodi, is little worn; 
indeed the specimen is so little abraded that the protocone is not yet 
properly connected to the protoconule. The pattern it presents is thus 
not typical, and it has been necessary to create a neotype which has 
been sectioned to show the simplified pattern consequent upon more 
advanced wear. The marked grooving of the mesostyle does not 
extend far down the tooth. It is also apparent in the type that the 
smooth and deep concavity of the two halves of the ectoloph is a 
feature of the early state of attrition and that further wear would 
show these to he a little more flattened. 

The upper teeth are intermediate in size between those of E. capensis 
and EH. quagga*, the breadth in Pm?—Pm* being about 27-30 mm., 
and in M,-M, about 25-29 mm. The halves of the ectoloph are 
concave with a slight flattening of the inner face, and the parastyle 
and mesostyle are prominent and somewhat abruptly marked off as 
in £. zebra*. The anterior face of the parastyle is obliquely flattened 
and commonly slightly grooved in the premolars. The protocone is 
elongate oval with a flattened or a “bilobed” inner wall. Its junction 
with the protoconule is not much in front of the middle of the proto- 
cone. The hypocone is small and has the appearance of a tendency 
towards isolation, as the hypoglyph is unusually deep in many 
specimens and there is also commonly an indentation in the wall of 
the medivallum between the hypocone and the metaconule. The 
caballine fold is usually present and may be quite strongly developed. 
The pli-protoconule is well developed and often deep, as also is the 
pli-postfossette. The pli-postfossette is usually clearly shown and 
secondary plications are common. The pli-hypostyle and_pli- 
protoloph are very variable in development and the former is 
sometimes well marked. 

The lower teeth are not certainly known, but from the normal size 
relationship between upper and lower teeth they should range from 
16-5 to 19 mm. for Pm? to Pm4 and 15-5 to 18 mm. for Mt to M?. The 
species, H. harrist Broom lies in the correct size group, and there is a 


454 Annals of the South African Museum. 


strong probability that HE. kuhni is the upper dentition of that 
species. 

In 1937 Broom and le Riche described and figured certain teeth 
which they referred to #. capensis. The Bothaville material has not 
been seen by the present writer, but the Sterkfontein specimens were 
amongst those lent to him by Broom. The lower jaw described by 
Broom and le Riche from Sterkfontein have the incisors and a second 
premolar present, and in size these exceed those of the living horses, 
but are otherwise not specifically identifiable. They might belong to 
E. capensis, EZ. kuhni, E. harrisi or E. plicatus. The upper palate 
and teeth from Sterkfontein were not found in association with the 
lower jaw. 

This upper jaw is well embedded in the typical rather hard cave 
matrix, and it has proved impossible to clean the specimen properly in 
the normal manner. The occlusal surface of some of these teeth had 
been ground down by Broom to expose the enamel pattern, and the 
pattern of the first right molar was figured in the paper cited (fig. 1, 
A, p. 770) and measurements given. When the present writer 
examined the material, however, it was realised that the ground 
surface was very oblique to the plane of normal wear with a con- 
sequent distortion and exaggeration of breadth. The whole specimen 
was accordingly carefully ground down parallel to the palate and the 
dentition exposed as it would appear in early wear. The surface of 
the specimen (Tvl. Mus., No. 682) with the patterns now exposed is 
shown, natural size, in fig. 24. The dimensions are: 


RPm2 RPm? RPm#4 RM} RM? RM? 
Breadth . 29:0 mm. 229mm. ?30mm. 27:5 mm. 227mm. ?26 mm. 
Length . 365mm. 295mm. 30mm. 305mm. 295mm. ?28 mm. 


The whole character of the teeth places the specimen in the species 
E. kuhni, of which it is an excellent example. 

The molar originally referred by van Hoepen (1930 a) to E. cawoodt, 
and later (1930 b) cited as the type of a new species H. lowwi, is a very 
typical tooth of EH. kuhni, as also are the other teeth ascribed to 
E. louwt. Haughton (1932) has already suggested that this species is 
a synonym of £. kuhnt, and this view is confirmed. 

The teeth which Dreyer (1931) refers to as “Equus kuhni simplex” 
are difficult to refer with certainty, but the “wholly unfossilised 
fourth premolar dug up at Glen Craig, Grahamstown”’, looks remark- 
ably like H. zebra*, while the one from Pniel is too small to be E£. 
kuhni and agrees excellently with little worn teeth of H. burchellu*. 
The partly fossilised first and second molars “dug up by Mrz. S. 


(‘TeuIS1Q) ‘oeZIs TBMyBNY “POOL oy} Surystjod puv Surpuri3 Aq posodxe 


3003 yooyo Joddn jo suseqqed joureue YyIM Puv XI1}eUT UT poppe 
‘moog wyny snnby—'ss “Ol 


ure ‘oABD UloJUOFY10}g Woy UoMToods porloyod JO MOTA [VFL]CA 


456 Annals of the South African Museum. 


Kinneard in Grahamstown”’, however, agree excellently both in size 
and character with #. kuhni and may certainly be assigned to that 
species. 


Fic. 24.—Hquus harrisi Broom. 
Top: Type left lower second molar and fourth premolar. (Original.) 
Centre: Referred right lower cheek teeth from Sterktontein Cave. (Original.) 
Below: Referred right lower fourth premolar from the Vaal River gravels at 
Pniel. (From Cooke.) 
All natural size. 


A Critical Revision of the Quaternary Perissodactyla. 457 


Equus harrist Broom. 


Equus harrist Broom 1928. Ann.S. Afr. Mus., xxii, pp. 441, fig. 2, B. 

Equus (Sterrohippus) harrisi (pars) Haughton 1932. Ann. §. Afr. Mus., 
XXvill, pp. 412-413. 

Equus capensis (pars) Cooke 1939. 8S. Afr. J. Sci., xxxvi, p. 413, 
fig. 2, A. 

(non Equus harrisi Dreyer 1931. ““New Fossil Mammals and Man”, 
pp. 23-25, pl. vi, figs. 18, 14; pl. vii, figs. 11, 12.) 

_ Lectotypes: Left lower Mand Pm,. M.M.K., No. 3939. (Fig. 24.) 

Locality and Horizon: “Diamond gravels of the Middle Terrace at 
the bend near Barkly West.” 


Measurements: 
LM, LPm, 
Breadth . : ‘ : : . 16-5 mm. 17-5 mm. 
Length . : : : : . 29-5 mm. 30-5 mm. 
Height (excluding roots) . : . 37:0 mm. 45-0 mm. 


The species is a good deal larger than EF. burchellii*, which it some- 
what resembles, and it is smaller than E. capensis. The type speci- 
mens are very much worn, and this fact makes specific identification 
with less worn specimens rather difficult. 

A feature of the species is its relatively greater antero-posterior 
compression compared with corresponding teeth in EH. capensis. In 
the premolars, the entoflexid lobe is simpler and smaller than in £. 
capensis. There is also a marked tendency in the premolars for the 
indentation between hypoconid and hypolophid to be very small or 
absent. The outer walls of the well-developed hypoconid and pro- 
toconid are flattened or even slightly concave inwards, the ectostylid 
fold in the hypoconid wall being small or lacking. There is a strongly 
developed entaconid of rounded or quadrate form. The metaconid 
is relatively large and elongate antero-posteriorly, while the meta- 
stylid is smaller and pear-shaped. The valley between metaconid 
and metastylid is bluntly pointed. 

Good material of this species is difficult to find, but an excellent 
series discovered by Broom in the hard limestone of the famous 
Sterkfontein Cave was given to the writer for identification and is 
figured here (fig. 24) for the first time.* It is unfortunately impossible 
to remove these teeth from their matrix, but the available dimensions 
are: 


RPm, RPm, RM, RM, 
Breadth  . 2 eb maim: 18 mm. 16-5 mm. 16 mm. 
Length ‘ . 33-0 mm. 32 mm. 28-5 mm. ?28 mm. 


* Also see Appendix. 
VOL. XXXI, PART 4. 38 


458 Annals of the South African Museum. 


A lower right fourth premolar from Pniel was identified and 
figured in error by the present writer (Cooke, 1939, p. 413, fig. 2) as 
belonging to EF. capensis. This tooth (Arch. Sur., No. 5) is now 
assigned to FE. harrisi and is refigured here. Its breadth is 17 mm., its 
length 30-5 mm. and its height 74 mm. 

The lower right fourth premolar (M.M.K., No. 4066), figured by 
Haughton (1932, p. 415, fig. 3, B) and referred by him to £. simplex 
van Hoepen, probably belongs to this species, as also may the 
associated right first molar (M.M.K., 4047). 


Referred Upper Teeth. 


In his original paper Broom (1928) refers to this species a right upper 
Pm‘ from the same locality. Considering the nature of this deposit, 
the fact of an associated occurrence is only of very slight value in con- 
necting the specimens. On a basis of size relationship between upper 
and lower teeth, the upper Pm* corresponding to the type lower tooth 
of this species should be about 28 or 29 mm. broad. The associated 
specimen referred by Broom has a transverse diameter of 31 mm. 
which, though not certainly outside the possible range, is rather on 
the large size. In view of these considerations the reference of this 
upper tooth to #. harrist cannot be regarded as satisfactory, and the 
tooth is therefore provisionally made the type of a new species Hquus 
broom described below. 

Van Hoepen (1930 a) referred this specimen to a new genus and 
species Sterrohippus robustus which he created on a second premolar. 
‘A second premolar cannot be regarded as forming adequate material 
for the type of a species, nor does there appear to be grounds for 
separating the genus from Equus. His specific name could not be 
retained as a species of this latter genus, as it is preoccupied by Equus 
robustus from the Pleistocene of Europe. Sterrohippus robustus is 
therefore regarded as incorrectly founded and invalid. 

The upper teeth of Equus harrisi must for the moment be regarded 
as unknown until they are found in a good association with recognis- 
able lower teeth. The rather poor lower teeth found in apparent 
association with upper teeth assigned to Equus kuhni Broom agree 
fairly well with those of H#. harrisi, but there is as yet insufficient 
material to warrant identification with that species. The upper 
teeth of HL. kuhni, however, are in precisely the right size group and 
it is very possible that EH. kuhni represents the upper dentition of 
E. harrist. The association of the two species in the Sterkfontein 
group of caves may also be significant. 


A Critical Revision of the Quaternary Perissodactyla. 459 


Equus plicatus (van Hoepen). 


Kolpohippus plicatus van Hoepen 1930. Pal. Nav. Nas. Mus. 


Bloemfontein, II, 1, pp. 8-10, fig. 10. 

Equus plicatus Dreyer 1931. “‘New Fossil Mammals and Man”, 
pp. 33-35. 

Equus plicatus Haughton 1932. Ann. 8. Afr. Mus., xxviii, p. 417. 


Type: Lower series Pm, to M, of the right side. Nas. Mus., No. 
C.425. (Fig. 25.) 

Locality: Tierfontein, near Port Allan, O.F-.S. 

Horizon: Not given in description; probably surface deposits. 

Measurements: The measurements cited in the type description do 
not agree with the text-figure, but it appears that the breadths given 
are total breadths and not just the width across the enamel. The 
dimensions given below give the breadth over the enamel, estimated 
by enlarging the illustration in van Hoepen’s paper to agree as well 
as possible with the various measurements he cites. The text-figure 
of the type series given here is the one derived in this way: 


RPm, RPm, RPm, RM, 
Breadth . . 17-5 mm. 17-5 mm. 15+ mm. 15-5 mm. 
Length . . 38:0 mm. 33-0 mm. 30 mm. 27-0 mm. 
Height . . 68-0 mm. 81-0 mm. 89 mm. 79-0 mm. 


+ very little worn. 


The lower teeth are larger than those of #. quagga* and smaller than 
those of E. capensis, agreeing in size with those of H. harrisi, which 
they otherwise closely resemble. The probable range of breadth in 
Pm, to Pm, is about 15-5 to 19 mm., and in M, to M, 14-5 to 17-5 mm. 
The outer walls of the protoconid and hypoconid are flattened or even 
concave inwards in the premolars; in the molars the protoconid may 
be convex. A strong ectostylid fold is present in the anterior part of 
the hypoconid wall in the premolars, but this fold is small or even 
absent in molars. The metaconid is rounded, and is separated from 
the bluntly pointed metastylid by a pointed groove. The entaconid 
is quadrate and smaller than the metaconid. The lobe of the ento- 
flexid is complicated in the premolars by strong—or at least well- 
marked—crimping of the inner wall of the hypoconid. The molars 
are much simpler than the premolars and cannot be easily distinguished 


from those of E. harrist. 
Van Hoepen’s reference of the type teeth to a new genus does not 


Fic. 25.—Equus plicatus (van Hoepen). 

Left: Crown view of type series of right lower cheek teeth. (After van 
Hoepen; modified.) 

Right: Crown view of series of lower left cheek teeth from damaged lower jaw 
from Koffiefontein, Orange Free State. This jaw is shown at the 
bottom of fig. 6. (Original.) 

Natural size. 


A Critical Revision of the Quaternary Perissodactyla. 461 


appear justifiable and even if the species is valid, it must be placed in 
Equus. 

A fine series of teeth in a damaged left lower jaw from Koffiefontein, 
O.F.S., may be referred to this species with considerable certainty. 
The jaw (M.M.K., No. 4845) is shown in fig. 6 in comparison with a 
skull of H. burchellii* and the enamel pattern of the teeth is shown in 
fig. 25. The inner side of the dentition is slightly damaged but the 
dimensions over the enamel are given below: 


LPm, LPm, LPm, LM, LM, 
Breadth : . L7 mm. 17-5 mm. 17-5 mm. 16:5 mm. 15:5 mm. 
Length ; . 38 mm. 33 mm. 33 mm. 29 mm. 29 mm. 


It would appear very probable that the differences between the 
dentitions ascribed to E. harrisi and to E. plicatus are not really of 
specific value. The type teeth of E. harrisi are very worn indeed, 
and it is difficult to assess the probable appearance of the teeth of this 
species in early wear. There can, however, be little doubt that teeth 
of L. plicatus in advanced wear would be indistinguishable from those 
assigned to #. harrist. 

The upper teeth are not known from a certain association. There 

is a strong probability that the upper dentition of a skull from the 
" same locality (Koffiefontein) as the lower jaw described above may 
be that of H. plicatus. This upper dentition has been described by 
Wells as E. fowler, and in the absence of a certain association the 
merging of these two species must remain an open question. 


Equus fowler: Wells. 


Equus fowleri Wells 1941. Trans. Roy. Soc. 8. Afr., xxvii, pp. 301- 
906, fig: 1, pl. Iv. 

Cotypes: “Portions of skull and isolated tooth catalogued as 
No. 555 in the collection of the Department of Anatomy, University of 
the Witwatersrand, Johannesburg.” (Figs. 6 and 26, A. B.) 

Locality: Koffiefontein, O.F.S. 

Horizon: “At a depth of about 18 inches in surface soil overlying 
shale.”’ 

Material: This comprises “the greater part of the right maxilla 
and palatine bone with Pm?—-M? in position and perfectly preserved, 
the right maxilla containing an unerupted I® and part of the left 
maxilla lacking teeth”. ‘‘The presence of the socket for C sug- 
gests that the individual is a male.” “The remains are those of an 
individual not quite fully grown.” 


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“Aqteooy od44 oy} Woy rvejour ysay soddn 4yoy od4400 Jo usz90y4ed UMOID “g 
“ad 44 Jo yy004 yooyo zoddn 4ystr Jo usr9yyed uMOID “Ww 
"say wajnof snnby—'9z "oy 


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A Critical Revision of the Quaternary Perissodactyla. 463 


Measurements: 

Skull: Length of upper jaw . f ‘ : . About 44-5 cm. 

Breadth across the first molars . : : . 16-5 em. 

Length of Pm?-M? : . 20-6 cm. 

Diastema between Pm? and I3 . Approximately 11 em, 

_ Teeth: 
em? Pm? Pm* M! M? M? 

Breadth . : 2 mime 3lmm. 31mm. 30mm. 29mm. 24mm. 
Length . : 44 mm. 309mm. 32mm. 32mm. 33mm. 32 mm. 
Length of  OEOROLS 13mm. 145mm. 15mm. 135mm. 15mm. 15 mm. 
Height (ex. roots) ~. 56 mm. 66mm. 70 mm. 64mm. 74mm. 76 mm. 


The following account is based on the type description, much being 
directly or indirectly quoted from Wells’ paper. 

The dimensions of the skull exceed those of many specimens of E. 
caballus* and considerably exceed those of the zebrine horses. The 
lateral aspect of the maxilla in its upper portion, below the naso- 
maxillary suture, shows a marked hollowing. This region thus 
presented a form seen in the asses and in the quagga group, but not in 
the caballine horses nor, to any marked degree, in the mountain zebra. 

The teeth are large, being intermediate in size between those of EH. 
capensis (EH. cawoodi) and E. kuhm, both of which they somewhat 
resemble. The halves of the ectoloph are deeply concave inwards, 
and curve rather sharply into the styles with a tendency for the 
mesostyle to overhang anteriorly. Both parastyle and mesostyle are 
prominent and massive, especially in the premolars, and are more or 
less conspicuously grooved. The protocone is elongate, and its 
connection with the protoloph is established very near its anterior 
extremity, so that the elongation appears to affect chiefly the posterior 
lobe. The medial wall of the protocone is sinuous, with an ill-defined 
median groove. In Pm? and Pm‘ the protocone has a remarkably 
oblique direction, and these two teeth in the type show an unusual 
prominence of the hypostyle.* The caballine fold is well developed 
and may be duplicated. The enamel of the fossettes is irregularly 
crimped and its secondary plications are numerous but relatively 
coarse. The pli-protoloph and pli-hypostyle are well developed in 
early wear. The pli-protoconule is unusually complex and even in 
M! is duplicated. The pli-postfossette is also rather complex, but the 
pli-prefossette is simpler, though small secondary folding may be 
present. 

Another tooth found half a mile distant from the skull fragments 
at a depth of five feet in similar soil has been referred to this species. 


* This is almost certainly a feature of early wear. 


464 Annals of the South African Museum. 


It is an upper left first molar and shows all the characteristics of the 
type, but is from a different individual, also young. The measure- 
ments of this tooth are: 


Breadth . : ; - ‘ . 30mm. 
Length : - : : ; . 32 mm. 
Height (excluding roots) : : . 65 mm. 


The pattern it presents is a little more complex in its secondary 
plication than that of the type (fig. 26, B). A section has been made 
through the tooth 15-20 mm. below the grinding surface, where the 
pattern is somewhat simplified but remains essentially the same 
(fig. 26, C.) 

In the McGregor Museum there occurs a right upper M? (M.M.K.., 
4342) also from Koffiefontein which Wells refers to his species (fig. 26, 
D). It is more worn than the corresponding tooth of the type and is 
a little smaller, its dimensions being: 


Breadth . 3 } : P . 28-5 mm. 
Length . : ; - . 29-5 mm. 
Height (excluding roots) . ‘ . 76 mm. 


The styles are narrower than in the type and are very indistinctly 
grooved, and the pli-hypostyle is single though remarkably large. 
These differences, however, are not outside the expected range of 
variation of such a species and the tooth is referred to E. fowlert. 

As has been stated above, the lower jaw which agrees so well with 
the type of van Hoepen’s #. plicatus, was found also at Koffiefontein, 
and this dentition is of a suitable size to correspond with £. fowlert. 
There is thus a good possibility that these names are synonymous, 
but such a decision must await confirmation from a better association. 

It also seems very likely that H. fowlert is merely a somewhat 
abnormal variant of E. kuhni, and that, in fact, BE. kuhni, E. harrist, 
E. plicatus and E. fowleri represent only a single species.* 


Equus sandwitht Haughton. 


Equus sandwithi Haughton 1932. Ann. 8. Afr. Mus., xxvii, pp. 
419-421, fig. 4. 
Cotypes: Upper right Pm? and Pm? and incomplete M°. 
Upper left Pm* and Pm+. 
Lower right M, and incomplete Pm. 
Lower left M, and M,. 
S.A. Mus., Nos. 6577 and 6578. (Fig. 27.) 
Locality: Usakos, South-West Africa. 


* Also see Appendix. 


A Critical Revision of the Quaternary Perissodactyla. 465 


yt ge aa 


L Meo LM, 


Fie. 27.—Equus sandwitht Haughton. Cotype upper and lower cheek teeth. 
Natural size. (Original.) 


VOL. XXXI, PART 4. 39 


466 Annals of the South African Museum. 


Horizon: In a bed of hard clay and sand 10 feet thick underlying 
8 feet of surface limestone. 


Measurements : 

Upper RPm? RPm?® RM RPm? LPm* 
Breadth ; . . 265mm. 28 (?) mm. 245 mm. 29mm. 32 mm. 
Length : : 2/4 poo at 285mm. 29(?)mm. 29mm. 29 mm. 
Height (ex. roots) . - 56mm. 54 mm. 67mm. 53mm. 57 mm. 
Length of protocone . “Timm. 10-5 mm. llmm. 9mm. 10mm. 

Lower RPm, RM, LM, LM, 
Breadth (Est.) : : . 16-17 mm. 15mm. 16mm. 11-5 mm. (14-5) 
Length ; : ; 30mm. 295mm. 29mm. 29mm. 
Height (ex. roots) . 70 mm. 78mm. 65 mm. 77 mm. 


Haughton’s measurements differ slightly from those given above, 
the explanation lying in the fact that the breadth quoted here is 
measured directly across the tooth and not along the rather sloping 
surface of wear. The tooth which Haughton describes as a lower left 
third molar is here regarded as a lower left second molar in very early 
wear. The very narrow enamel surface is a feature of early wear, 
and the crown is 3 mm. wider half an inch below its present surface. 

The upper teeth of this species resemble those of Equus zebra* but 
are a good deal larger and present an even simpler enamel pattern. 
The halves of the ectoloph are flattened or even bulge outwards near 
their centres. The parastyle and mesostyle are strongly developed 
and well demarcated. The mesostyle is rather narrow, and the 
parastyle has an oblique flattened face which may be grooved. The 
protocone is abnormally small, being proportionally even shorter 
than in £. zebra*. This shortness is due to the very small develop- 
ment of the anterior prolongation. The fossettes are large and 
lunate. They are almost devoid of folding, only the pli-protoconule 
being distinct while the other folds are almost indistinguishable. The 
pli-caballin is absent. The hypoglyph is shallow. The protoconule 
shows marked antero-internal flattening. 

The lower teeth are unusual in shape, as they present an anterior 
face which is oblique to the axis of the tooth instead of directly 
transverse to it. This is due to the posterior deflection of the point 
of the parastylid and its anterior face. The walls of the protoconid 
and hypoconid are rounded or slightly flattened and the ectostylid 
fold is inconspicuous or absent. The entaconid is moderately small in 
the molars. The metaconid and metastylid are normally large and 
pear-shaped, but their attitude may be distorted by the relative 
compression of the interior side of the tooth. The anterior part of the 
entoflexid tends to be slightly or moderately folded. 


A Critical Revision of the Quaternary Perissodactyla. 467 


The ratio of size of molars and premolars is rather unusual, as the 
molars appear relatively to be abnormally small and narrow both in 
upper and lower dentitions. The type LPm*‘ owes its unusual width 
at least in part to the abnormal projection of the mesostyle, but it 
appears that the expected range in breadth would be 28-33 mm. in 
Pm? and Pm* and 26-29 mm. in M! and M?, with the anterior and 
posterior teeth notably narrower. The same applies to the lower 
teeth, in which the range in Pm, and Pin, may be 15-18 mm. and in 
M, and M, 13-17 mm. 


Referred Material. 


Haughton (1932) assigned to H. sandwithi two lower teeth from the 
Vaal River gravels at Sheppard Island. These teeth were discovered 
by van Riet Lowe but were lost in the library fire at the University of 
the Witwatersrand in 1931. 


LPm, 


RP? 


RPm4 


Fic. 28.—Equus sandwithi Haughton. Referred upper and lower cheek teeth 
from the Kromdraai Cave. Natural size. (Original.) 


Amongst the material collected by Broom in the Kromdraai Cave 
deposit there occur several teeth which appear to belong to this 
species. Most of the teeth, both upper and lower, are damaged and 
difficult to clean, but a selection of the better ones 1s shown in fig. 28. 


Equus poweri Cooke. 
Equus powert Cooke 1939. Q Afr. J. Sci., XXXVI, PP. paul 
fig. 1. 


468 Annals of the South African Museum. 


Type: Upper right first molar. Arch. Sur., No.3. (Fig. 29). 
Locality: Pniel, Vaal River. 

Horizon: ?% Younger gravels of Vaal River. 

Neotype: Upper left first molar. Arch. Sur., No. 213. (Fig. 29.) 
Locality: 35-foot shaft, Lot 197, Windsorton. 

Horizon: Younger gravels of Vaal River. 


Fic. 29.—Equus poweri Cooke. 
Left: Type right upper first molar. Outer and crown views. 
Right: Neotype left upper first molar. Outer and crown views. 
Natural size. (Original.) 


Measurements: 
Type RM? Neotype LM? 
Breadth . ‘ . 27 mm. 26 mm. 
Length . - ‘ 24 mm. 23 mm. 
Height . : : 44 mm. 64 mm. 


This species resembles Equus sandwitht, but appears to be very 
slightly smaller and to differ in several particulars. The halves of the 


A Critical Revision of the Quaternary Perissodactyla. 469 


ectoloph are moderately flattened or bulge outwards near their 
centres, and the parastyle and mesostyle are well demarcated. 
Though prominent, the mesostyle is not nearly as well developed as 
in EL. sandwith. The protocone is very small, but the anterior lobe 
is proportionately larger than in E. sandwithi and more like E. zebra* 
in this respect. The fossette are large and simple, the post-fossette 
being subrectangular in shape and not lunate asin H. sandwitht. The 
pli-protoconule is small and the hyposiyle and protoloph folds are 
present only as small notches. The medivallum is small and narrow 
and devoid of any trace of a caballine fold. The protoconule shows a 
well-rounded antero-internal face and is not flattened as in #. sand- 
with. The hypoglyph is very shallow and may disappear altogether 
towards the base of the tooth. 

The estimated dimensions fall a millimetre below those of E. 
sanduitht. 


Equus broom sp. nov. 


Equus harrisi (pars) Broom 1928. Ann. 8. Afr. Mus., xxii, pp. 442- 
443, fig. 2 (B,). 
Type: Upper right fourth premolar. M.M.K., No. 3939 c. (Fig. 
30.) 
Locality: The Bend, near Barkly West. 
Horizon: “Middle Terrace” of the Vall River gravels. 
Measuremenis : 


RPm* 
Breadth . : : : -)  olemm. 
Length 2 : : : - o2mm. 
Height ‘ : : ‘ . 62mm. 
Length of protocone . ‘ : 9 mm. 


This tooth, which is badly battered, was assigned by Broom to the 
species E. harrisi on account of the association of the specimen with 
the two lower type teeth of that species. This association is not a 
good one in view of the very disturbed nature of the deposit, and the 
upper tooth seems somewhat large to be correctly associated with the 
lowers. Itis therefore given a new name here until certain associations 
settle the problems of the relations between the various species named 
on upper and on lower teeth. 

The resemblance of this species to Equus sandwithi is strong, almost 
the only marked difference, apart from the greater size, being the shape 
of the ectoloph, which is smoothly concave in both halves and is only 
slightly overhung by the parastyle. The protocone is very short and 


470 Annals of the South African Museum. 


unusually rounded. The hypocone projects into the medivallum as 
a slight bulge but the hypoglyph is small. The fossettes show more 
folding than is the case in H#. sandwitht, the pli-protoloph, pli- 
postfossette and pli-protoconule being distinct though small, and 


Fie. 30.—Equus broomi sp. nov. Outer lateral and crown views of the 
type upper right fourth premolar of Equus broomi sp. nov.; formerly 
referred to Equus harrisi Broom. Natural size. (Original.) 


there is a certain amount of minor folding in the posterior wall of the 
prefossette. | 

The specimen on which van Hoepen created the new genus Sterro- 
hippus and species robustus might belong to this species, but van 
Hoepen’s specimen is a second premolar which cannot be regarded 
as forming adequate material for specific identification. There is no 
reason to support the generic status of Sterrohippus, which is indis- 
tinguishable from many species of Equus, and the specific name Hquus 


A Critical Revision of the Quaternary Perissodactyla. 471 


robustus is preoccupied and cannot be retained. Sterrohippus robustus 
is therefore regarded as invalid. 


Incertae sedis. 
Equus simplex van Hoepen 1930. Pal. Nav. Nas. Mus. Bloemfontein, 
SI p25; fig. 6. 


Type: Upper-right first molar. (Nas. Mus., No. 284). 
Locality: Koffiefontein, O.F-.S. 
Horizon: % Surface. 


Measurements: 
RM1 
‘ Breadth . , . 380mm. 
Length . : - ag) naan, 
Height . : - _o9 mm. 


The type specimen has not been seen by the writer, and it is not 
possible from the drawing to assign the tooth with certainty to any 
particular species. If the dimensions given in the text of the type 
description are correct, the tooth may belong to FE. capensis as 
suggested by Haughton (1932). Otherwise from the text-figure it 
resembles #. kuhna more closely than H. capensis, so that the position 
(or the validity) of this species is uncertain. 


GuIDE TO EQUINE SPECIES. 


Incisors. 


A. Very broad mandibular symphysis with large, anteriorly flattened 
first and second incisors and reduced third incisors. (Upper 
incisors unknown.) 

Eurygnathohippus cornelianus v. Hoep. 


B. Incisors of the normal (i.e. Equus caballus*) type possessing the 
cup or “mark” in the upper jaw but lacking it in the lower 
jaw. 

Equus burchellic* (Gray). 
C. Incisors of the “normal” type with the cup or “mark” present at 
least in the first and second incisors of both jaws. 
2 Hquus quagga* Gmelin. 
Equus zebra* Linn. 
? Hquus capensis Broom. 
? Equus kuhn Broom. 
2 Equus fowlert Wells. 


472 Annals of the South African Museum. 


D. Incisors unknown. 
Equus harrist Broom. 
Equus plicatus (v. Hoep.). 
Equus sandwitht Htn. 
Equus poweri Cooke. 
Equus broom: Cooke. 
(Equus simplex v. Hoep.) 
Notohipparion namaquense Htn. 
Stylohipparion steytlert (v. Hoep.). 


Upper Cheek Teeth. 


A. Protocone isolated; parastyle anteriorly flanged. 
Stylohipparion steytlert (v. Hoep.). 
(Pm. ? 19-24 mm. M. 18-23 mm.) * 


B. Protocone attached. 
1. Breadth less than 27 mm. 
(a) Ectoloph halves concave; parastyle obliquely flattened 
anteriorly. 
Equus burchellu* (Gray). 
(Pm. 22-26 mm. M. 21-25 mm.) 


(b) Ectoloph halves flattened. 
Equus zebra* Linn. 
(Pm. 22-27 mm. M. 21-26 mm.) 
Equus quagga* Gmelin. 
(Pm. 22-27 mm. M. 21-26 mm.) 


2. Breadth greater than 27 mm. 
(a) Ectoloph halves concave; parastyle usually flattened. 

Equus capensis Broom. 
(Pm. 29-35 mm. M. 27-33 mm.) 
Equus kuhni Broom. 
(Pm. 27-30 mm. M. 26-29 mm.) 
Equus broom: Cooke. 
(Pm. 28-33 mm. M. 27-32 mm.), 
Equus fowlert Wells. 
(Pm. 29-33 mm. M. 28-32 mm.) 


(6) Kctoloph halves flattened; parastyle prominent. 
Some #. kuhni premolars. 


* These dimensions are range of breadth. 


A Critical Revision of the Quaternary Perissodactyla. 473 


Equus sandwithi Htn. 

(Pm. 28-33 mm. M. 26-29 mm.) 
Equus powert Cooke. 

(Pm. ? 26-29 mm. M. 25-28 mm.) 


. Position uncertain. 


Equus simplex v. Hoep. 


. Upper cheek teeth unknown. 

Equus plicatus (v. Hoep.). 

Equus harrist Broom. 

Notohipparion namaquense Htn. 
Eurygnathohippus cornelianus v. Hoep. 


Lower Cheek Teeth. 


. External ectostylid column present. 
1. Strong protoconid fold or pillar; metaconid and metastylid 
strong. 
Notohipparion namaquense Htn. 
(Pm. 15-16 mm. M. 14-17 mm.) 


2. Protoconid fold weak or lacking; metaconid and metastylid 
small. 
Stylohipparion steytleri (v. Hoep.). 
(Pm. 11-15 mm. M. 10-13 mm.) 


. No external ectostylid column. 

1. Face of parastylid oblique; metaconid and metastylid strong. 
Equus sandwitht Htn. 
(Pm. 15-18 mm. M. 13-17 mm.) 


2. Face of parastylid normal. 
(a) Breadth less than 15 mm. 
Equus zebra* Linn. 
(Pm. 12-15 mm. M. 11-14 mm.) 
Equus burchelli* (Gray). 
(Pm. 12-15 mm. M. 11-14 mm.) 
Equus quagga* Gmeiin. 
(Pm. 12-15 mm. M. 11-14 mm.) 
(Some molars of E. plicatus resemble LE. quagga*.) 


(b) Breadth greater than 15 mm. 
Equus harrisi Broom. 
(Pm. 17-20 mm. M. 16-19 mm.) 


474 Annals of the South African Museum. 


Equus plicatus (v. Hoep.). 

(Pm. 15:5-19 mm. M. 14:5-17-5 mm.). 
Equus capensis Broom. 

(Pm. 18:5-22 mm. M. 17-5-21 mm.) 


C. Lower dentitions unknown. 
Eurygnathohippus cornelianus v. Hoep. 
Equus kuhni Broom. 

Equus fowlert Wells. 
Equus broom: Cooke. 
Equus powert Cooke. 


REFERENCES. 


ALLEN, G. N., 1939. ‘““A Checklist of African Mammals”, Bull. Mus. of Compar. 
Zool. Harvard, vol. lxxxiii, pp. 1-763. 

Brcx, R., 1906. “Mastodon in the Pleistocene of South Africa”, Geol. Mag., 
Decade V, vol. iii, pp. 49-50. 

Broom, R., 1909. “‘On the Evidence of a Large Horse recently Extinct in South 
Africa”’, Ann. S. Afr. Mus., vol. vii, pp. 281-282. 

——, 1913 a. “Note on Equus capensis’’, Bull. Amer. Mus. Nat. Hist., vol. xxxii, 
pp. 437-439. 

——, 19136. ‘Man Contemporaneous with Extinct Animals”, Ann. 8. Afr. Mus., 
vol. xii, pp. 13-16. 

——, 1928. ‘On Some New Mammals from the Diamond Gravels of the Kimberley 
District”’, Ann. S. Afr. Mus., vol. xxii, pp. 439-444. 

—, 1937. ‘‘New Pleistocene Mammals from Limestone Caves of the Transvaal’’, 
S. Afr. J. Sci., vol. xxxiii, pp. 750-768. 

Broom, R., and Lz Ricuez, H., 1937. ‘The Dentition of Equus capensis Broom”’, 
8. Afr. J. Sci., vol. xxxiii, pp. 769-770. 

Cuuss, E. C., 1907. “List of Vertebrate Remains” in “On an African Occurrence 
of Fossil Mammalia associated with Stone Implements’’, by F. P. Mennel and 
KE. C. Chubb, Geol. Mag., Decade V, vol. iv, pp. 447-448. 

Cooxg, H. B. S., 1939. “On a Collection of Fossil Mammalian Remains from the 
Vaal River Gravels at Pniel”’, S. Afr. J. Sci., vol. xxxvi, pp. 412-416. 

——, 194la. “A Preliminary Account of the Wonderwerk Cave, Kuruman 
District. Section II.: The Fossil Remains”, S. Afr. J. Sci., vol. xxxvii, pp. 
303-311. 

—, 19416. “A Preliminary Survey of the Quaternary Period in Southern 
Africa”, Bur. of Archaeology, Arch., Ser. iv, pp. 1-59. 

——,1948. ‘Cranial and Dental Characters ofthe Recent South African Equidae’’, 
S. Afr. J. Sci., vol. xl, pp. 254-257. 

(See also WELLS, CooKE and Maan, 1942; Matan and Cooks, 1941.) 


A Critical Revision of the Quaternary Perissodactyla. 475 


Dreyer, T. F., in Dreyer, T. F., and Lyuz, A., 1931. ‘New Fossil Mammals and 
Man from South Africa”, Dept. Zool. Grey. Univ. College, Bloemfontein. 
Fraas, E., 1907. » Pleistocane Fauna aus dem Diamantseifen van Siid-Afrika”’, 

Zeit. d. Deutsch. geol. Gesell., vol. lix, pp. 232-243. 

GIDLEY, J . W., 1901. ‘“‘Tooth Characters and Revision of the North American 
Species of the Genus Hquus”’, Bull. Amer. Mus. Nat. Hist., vol. xiv, pp. 91-141. 

Grecory, W. K., 1934. “A Half-Century of Trituberculy: the Cope-Osborn Theory 
of Dental Evolution”’’, Proc. Amer. Phil. Soc., vol. Ixxiii, No. 4, pp. 169-317. 

HaveutTon, 8. H., 1932. “The Fossil Equidae of South Africa”’, Ann. 8. Afr. Mus., 
vol. xxviii, pp. 407-427 (February 1932). 

Horwoop, A. T., 1926. “Fossil Mammalia” in “Geology and Palaeontology of 
the Kaiso Bone Beds’’, Occ. Paper No. 2, Geol. Survey of Uganda. 

—, 1928. ‘‘Mammalia” in Rhodesian Man and Associated Remains, London: 
British Mus. (Natural History), pp. 70-73. 

Le Ricus, H., 1937. See Broom, R., 1937. 

Linnaeus, C., 1758. Systema Naturae, 10th Edition. 

Lowe, C. vAN Rizt, 1937. See S6HNGE, VissER and Lowe, 1937. 

Maay, B. D., and Cooks, H. B.S.,1941, ‘‘A Preliminary Account of the Wonder- 
werk Cave, Kuruman District”, 8. Afr. J. Sci., vol. xxxvii, pp. 300-312. 

Matay, B. D., 1943. See Wetts, L. H., 1943. 

Ossorn, H. F., 1907. Evolution of Mammalian Molar Teeth. New York: 
Macmillan & Co. 

(See also Scorr and Ossory, 1890). 

Owen, R., 1869. “Description of the Cavernof Bruniquel and its Organic Contents. 
Part II. Equine Remains”, Phil. Trans. Roy. Soc. London, vol. clix, pp. 
535-557. 

Romer, A. S., 1933. Vertebrate Paleontology. Chicago: University of Chicago 
Press. 

Scort, W. B., 1907. ‘‘Mammalian Remains from the Coast of Zululand’’, Third 
and Final Report, Geol. Survey Natal and Zululand, pp. 251-262. 

Scott, W. B., and Ossorn, H. F., 1890. ‘“‘Preliminary Account of the Fossil 
Mammals from the White River and Loop Fork Formation contained in the 
Museum of Comparative Zoology. The Perissodactyla”, by H. F. Osborn, 
Bull. Mus. Comp. Zool., vol. xx, No. 3. 

Szriey, H. G., 1891. “On Bubalus bainii (Seeley)”, Geol. Mag., vol. viii, pp. 
199-201. 

Suaprro, M. M. J., 1943. “Fossil Mammalian Remains from Bankies, Kroonstad 
District, O.F.S.”, S. Afr. J. Sci., vol. xxxix, pp. 176-181. 

Suaw, J. C. M., 1938 a. “The Teeth of the South African Fossil Pig (Notochoerus 
capensis syn. meadowsi) and their Geological Significance”, Trans. Roy. Soc. 
S. Afr., vol. xxvi, pp. 25-37. 

——, 1938. b.. “Growth, Changes and Variations in Warthog Third Molars and 
their Palaeontological Importance”, Trans. Roy. Soc. S. Afr., vol. xxvii, 
pp. 51-94. 

Sounce, P. G., Visser, D. J. L., and Lows, C. van Riet, 1937. “Geology and 
Archaeology of the Vaal River Basin”, Geol. Mem. 35, Geol. Survey of the 
Union of S. Afr. 

Srirton, R.A., 1941. “Development of Characters in Horse Teeth and the Dental 
Nomenclature”, Journ. of Mammalogy, vol. xxit, pp- 434-446. 


476 Annals of the South African Museum. 


van Horpen, E. C. N.,1930a. ‘‘Vrystaatse Fossiele Perde’’, Pal. Nav. Nas. Mus. 
Bloemfontein, II, pp. 1-11. 

——, 1930 b. ‘‘Fossiele Perde van Cornelia, O.V.S.”, Pal. Nav. Nas. Mus. 
Bloemfontein, II, pp. 13-24. 

—, 1932. ‘Die Stamlyn van die Sebras”’, Pal. Nav. Nas. Mus. Bloemfontein, IT, 
pp. 25-37 (September 1932). 

—, 1940. “Oor die tande van die Equinae: 1. Die snytande van die 
Onderkaak”’, Tydskrif vir Wetenskap en Kuns, vol. i, pp. 101-114. 

Visser, D. J. L., 1937. See S6uncE, P. G., 1937. 

WELLS, L. H., 1941. ‘A Fossil Horse from Koffiefontein, O.F.S.”, Trans. Roy. 
Soc. 8. Afr., vol. xxviii, pp. 301-306, pl. lv. 

WELLS, L. H., Cooxsn, H. B. S., and Maan, B. D., 1942. ‘*‘The Associated Fauna 
and Culture of the Vlakkraal Thermal Springs, O.F.S.’’, Trans. Roy. Soc. 8. Afr., 
vol. xxix, pp. 203-233. 

WELLS, L. H., and Maran, B. D., 1943. “A Further Report on the Wonderwerk 
Cave, Kuruman. Section I: Archaeology, by B. D. Malan. Section JI: 
Fauna, by L. H. Wells”, 8S. Afr. J. Sci., vol. xl, pp. 258-270. 

ZITTEL, K. A. von, 1925. Text-book of Palaeontology. Vol. III. ““Mammalia”’, 
ed. by Sir A. Smith Woodward. London: Macmillan & Co. 


A Critical Revision of the Quaternary Perissodactyla. 477 


APPENDIX 
(With Fig. 31.) 


In the five years which have elapsed since this account of the Perisso- 
dactyla was written, certain additional information has come to light 
which, while it does not involve any important alteration of the views 
already expressed, nevertheless requires to be added in order to bring 
the work reasonably up to date. 

Firstly, the description of the fossil mammals of the Vaal River 
deposits, mentioned in the Introduction to the present account, has 
appeared as Memoir 35 (III) of the Geological Survey of the Union of 
South Africa and it contains a certain amount of stratigraphic informa- 
tion on the distribution of the Equidae in those deposits (Cooke, 
1949 6). In addition, revisions of the Suina (Cooke, 1949 a) and of the 
Proboscidea (Cooke, 1947) have been published as companion studies 
to the present one. 

Secondly, a perissodactyl family new to this region has been 
recorded through the discovery of an undoubted Chalicothere in the 
cave breccia of the Makapan valley (George, 1950). The type isa left 
upper second molar and there are also a number of other cheek teeth 
and an ungual phalanx. The type molar and the phalanx are 
illustrated here (fig. 31). Chalicothere remains have previously 
been reported from the Kaiso beds of Uganda (Andrews, 1923) and 
from south Serengeti in Tanganyika (Dietrich, 1942), but the Makapan 
specimens appear distinct and were described as a new species 
tentatively placed in Metaschizotherum as M. transvaalensis. The 
resemblance to the Tanganyika species M. henmgi is fairly close 
and it is considered by Miss George that both species certainly belong 
to the same genus, whether or not this is really Metaschizotherium. 

Thirdly, a new species of fossil equine has been erected by Broom 
(1948) based on material from a cave breccia in the Sterkfontein area 
about half a mile south-west of the site known as “Bolt’s workings”’. 
The type of this species, Equus zietsmani, comprises an imperfect 
lower jaw with the incisors, most of the cheek teeth of the right side 
and some of those of the left side, and part of the left maxilla. The 
lower series from P, to M, can be well reconstructed and is figured by 


478 Annals of the South African Museum. 


Broom, as also is the damaged upper series from P* to M (the latter 
tooth being a mere fragment). The lower series agrees very well with 
that shown in fig. 24 of the present text and referred to E. harrist. The 
upper series resembles that illustrated in fig. 23, which was referred 
to E. kuhni, and also shows features suggestive of #. fowlerr. In view 
of these resemblances and especially as the teeth shown in figs. 23 and 
24 were also from the Sterkfontein group of breccias, it is likely that 
only a single species is represented by the three lots of specimens. It 
remains to be considered what that species is. 


Fic. 31.—Metaschizotherium (?) transvaalensis George. 


Top left: Anterior view of ungual phalanx. 
Lower left: Side view of ungual phalanx. 
Right: Crown view of type left upper second molar. 
All natural size. (From George.) 


It has already been suggested in the present revision (p. 464) that 
Equus fowlert and EH. kuhni may be synonyms and that the corre- 
sponding lower dentitions may be represented by F#. plicatus and 
E. harrist. This idea lacked confirmation due to the absence of 
associated upper and lower dentitions but Broom’s new material goes 
a long way towards correcting this deficiency and greatly strengthens 
the probability that all five species are identical. It may now be 
considered that EF. fowleri, E. plicatus, E. harrisi and E. zietsmani are 
synonyms of Equus kuhni. 

Some additional confirmation of the above conclusion can be 
obtained from a study of the dental characters of the living Grevy’s 


A Critical Revision of the Quaternary Perissodactyla. 479 


Zebra of Kast Africa. It was suggested to the writer by Dr. L. S. B. 
Leakey that the South African Equus kuhni was, in fact, Equus 
grevyt. Through the courtesy of Dr. Leakey the writer had the 
opportunity, while in Hast Africa, of studying a small collection of 


Grevy skulls and it is clear that there is a close resemblance between 


the teeth of the living animal and the fossil material referred to 
E. kuhm. Individual teeth, but not whole dentitions, exhibited 
characters recalling H. fowler, EL. plicatus and EH. harrisi and the 
suggested identity of the four fossil species thus receives a measure of 
confirmation. There can be no doubt that the fossil HL. kuhni (and 
its presumed synonyms) is closely related to H. grevyi and the relatively 
small differences which have been noted could be dismissed as due to 
individual variation. For the present, however, it is felt that it will 
be better to retain the designation EH. kuhni for the fossil material 
until additional skeletal material is available to support the absorption 
of LE. kuhni into EF. grevyt. 


REFERENCES. 


AnpreEws, ©. W., 1923. In Nature, 10th November 1923. 

Broom, R., 1948. ‘‘Some South African Pliocene and Pleistocene Mammals”’, 
Ann. Tvl. Mus., vol. xxi, pp. 23-25. 

Cooxs, H. B. S., 1947. ‘‘ Variation in third molars of the living African Elephant 
and a Critical Revision of the Fossil Proboscidea of Southern Africa”’, Amer. 
J. Sci., vol. cexlv, pp. 434-457, 492-517. 

——, 1949a. “The Fossil Suina of South Africa’’, Trans. Roy. Soc. 8. Afr., 
vol. xxxii, pp. 1-44. 

——, 1949. ‘“‘Fossil Mammals of the Vaal River Deposits’’, Geol. Mem. 35 (IIT), 
Geol. Survey of the Union of 8. Afr., pp. 1-109. 

Dierricu, W. O., 1942. In Palaeontographica, Band xciv, Abt. A, 94. 

Gzrorcs, M., 1950. ‘‘A Chalicothere from the Limeworks Quarry of the Makapan 
Valley, Potgietersrust District”, S. Afr. J. Sci., vol. xlvi, pp. 241-242. 


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